Invasive, naturalized and casual alien plants in southern Africa

Invasive, naturalized and casual alien plants in southern Africa: a sum-mary based on the Southern African Plant Invaders Atlas (SAPIA)

L. HENDERSON*

Keywords: biomes, casual alien plants, invasive plants, Lesotho, naturalized plants, roadside surveys, SAPIA mapping project, South Africa, Swaziland

ABSTRACT

The primary objective of this publication is to provide an overview of the species identity, invasion status, geographical extent, and abundance of alien plants in South Africa, Swaziland and Lesotho, based on fi eld records from 1979 to the end of 2000. The dataset is all the species records for the study area in the Southern African Plant Invaders Atlas (SAPIA) database during this time period. A total of 548 naturalized and casual alien plant species were catalogued and invasion was recorded almost throughout the study area. Most invasion, in terms of both species numbers and total species abundance, was recorded along the southern, southwestern and eastern coastal belts and in the adjacent interior. This area includes the whole of the Fynbos and Forest Biomes, and the moister eastern parts of the Grassland and Savanna Biomes. This study reinforces previous studies that the Fynbos Biome is the most extensively invaded vegetation type in South Africa but it also shows that parts of Savanna and Grassland are as heavily invaded as parts of the Fynbos. The Fabaceae is prominent in all biomes and Acacia with 17 listed species, accounts for a very large proportion of all invasion. Acacia mearnsii was by far the most prominent invasive species in the study area, followed by A. saligna, Lantana camara, A. cyclops, Opuntia fi cus-indica, Solanum mauritianum, Populus alba/×canescens, Melia azedarach, A. dealbata and species of Prosopis.

INTRODUCTION

History of roadside surveys in South Africa

Roadside surveys of invasive plants in South Africa were pioneered by Henderson and Musil (née Duggan) starting in 1979 in the central Transvaal, now Gauteng (Wells, Duggan & Henderson 1980), with the remainder of the Transvaal surveyed in 1982 and 1983 (Henderson & Musil 1984). Surveys of the rest of South Africa were conducted by Henderson from 1986, starting with Natal (Henderson 1989), followed by the Orange Free State (Henderson 1991a), northern Cape (Henderson 1991b), eastern Cape (Henderson 1992), western and central Cape (completed in 1993 but unpublished), and southern and southwestern Cape (Henderson 1998a).

All terminology used in this paper relating to invasive plants such as ‘alien’, ‘invasive’, ‘naturalized’, ‘casual alien’, ‘weed’ and ‘environmental weed’ conforms, as far as possible, to the defi nitions provided by Richardson et al. (2000) and Pyšek et al. (2004). The method used in these surveys was designed initially to make use of otherwise unproductive travelling time whilst engaged in other research projects. The method was refi ned as the surveys progressed until a standardized method was developed (see Henderson 1992, 1998a). The presence and abundance of all alien trees, large shrubs and con-spicuous climbers which appeared to be naturalized or occurring outside of cultivation were recorded for each veld type category, habitat type (roadsides and adjoining veld, and streambanks) and quarter-degree/fi fteen minute square traversed by road.

Recordings of species on roadsides and in the adja-cent veld were made from a moving vehicle along road

transects of between fi ve and 10 km long. Recordings of streambank species were made at virtually all water-course crossings on the survey route.

The Southern African Plant Invaders Atlas mapping project (SAPIA)

The Southern African Plant Invaders Atlas (SAPIA) is a mapping project, launched in January 1994, to collate information on the distribution, abundance and habitat types of invasive and naturalized alien plants in southern Africa (Henderson 1998b). The fi rst phase of SAPIA, involving volunteer participants, was scheduled for a fi ve-year period, ending in December 1998. The atlas region covered South Africa, Lesotho and Swaziland. Information was recorded on two standardized atlas sheets, with slightly different species lists, covering the western and eastern halves of the atlas region. One hun-dred plant taxa were listed on each sheet, with a com-bined total of 161 species. A pocket fi eld guide was com-piled to help with the identifi cation of all listed species (Henderson 1995).

SAPIA database

A computerized SAPIA database was created by incorporating all Henderson survey data (± 23 000 records) and SAPIA phase one project data (± 20 000 records). The SAPIA project continued on an ad hoc basis and by the end of 2000 a total of ± 48 000 records had been accumulated. Thereafter, the SAPIA initiative dwindled due to lack of funding. Only 10 000 records were added in the fi ve year period from 2001 to the end of 2005. The SAPIA project was revived in 2006 with funding from the Department of Water Affairs and Forestry’s Working for Water Programme. The SAPIA database has been computerized using Microsoft Access and is housed at the Plant Protection Research Institute in Pretoria.

* Agricultural Research Council: Plant Protection Research Institute, c/o SANBI, Private Bag X101, 0001 Pretoria.E-mail: [email protected]. received: 2006-10-06.

Bothalia 37,2: 215–248 (2007)

216 Bothalia 37,2 (2007)

Objectives of this study

• To provide an overview of the species identity, inva-sion status, geographical extent, and abundance of alien plants in South Africa, Swaziland and Lesotho, based on fi eld records from 1979 to the end of 2000.

• To highlight the most prominent invaders in the region as a whole, in each of the biomes, and in riparian and wetland habitats.

• To compare invasion and provide species profi les for each of the biomes.

METHODS

Sampling method

The dataset for this study is all the species records for South Africa, Swaziland and Lesotho in the SAPIA data-base collected from 1979 until the end of 2000 (± 48 000 records). During this period a concerted effort was made to gather as much data from as wide an area as possi-ble. The information gathered is the best available data concerning the extent of invasion and species composi-tion, at least of the larger trees, shrubs and conspicuous climbers, in the study area over this time period.

The SAPIA dataset was subdivided on a quarter-degree square (QDS) basis into six datasets representing the biomes of southern Africa. According to Rutherford (1997) there are seven biomes in southern Africa: Savanna, Fynbos, Forest, Grassland, Nama-Karoo, Succulent Karoo and Desert. The Forest Biome in southern Africa is min-iscule, only occurring in the Knysna area. However, if all the forest patches elsewhere are included, its area increases several-fold (Rutherford 1997). In this study Forest refers to the Forest Biome and also forest habitats within the Savanna, Fynbos and Grassland Biomes. The Desert Biome occurs almost exclusively in Namibia, except for a very small patch along the Orange River bordering on South Africa that has been excluded from this study.

Data treatment

Abundance

Species abundance ratings in the SAPIA database are qualitative estimates. Table 1 shows the abundance rat-ings used in the SAPIA database and the equivalent rat-ing used in Henderson surveys. For the purposes of this study, species abundance ratings were converted to a numerical value as done in previous surveys (Henderson 1998a) and each abundance rating was expressed in numbers of individuals or groups per 10 km transect/recording (Table 1).

Prominence

A similar formula was used in this study to calcu-late prominence as in previous studies by Henderson (1998a). The prominence value of a species x in category y (biome or study area) was calculated as follows:

total abundance of species x in category y × 100sum of the abundances of all species in category y

prominence value

total species records of species x in category y × 100sum of the records of all species in category y

The highest prominence values in a given category which add up to ± 160 points out of a total of 200 are printed in bold in Appendices 1–3. The cut-off point is arbitrary but represents the upper 80% of the summed prominence values.

RESULTS

A total of 548 naturalized and casual alien plant spe-cies were catalogued in the SAPIA database for South Africa, Swaziland and Lesotho from 1979 to the end of 2000 (Appendix 4). At least 119, mainly herbaceous, taxa are considered to have been under-recorded and

TABLE 1.—Abundance ratings used in Henderson surveys, SAPIA and this study

Henderson surveys SAPIA This studyRating Roadsides and veld No.* Streambanks Rating All habitats† All habitats#

9 A virtually continuous, almost pure stand 1 000+ Any number, with cover more than 75% of the reference area

7 Very abundant 1 000

8 The commonest species in a generally con-tinuous tree or shrub layer

500–999 Any number, with 50–75% cover 6 Very abundant 1 000

7 Less abundant than above but > 20 individu-als or groups per km

200–499 Any number, with 25–50% cover 5 Abundant 200

6 10–20 individuals or groups per km 100–199 Any number with 5–25% cover 4 Abundant 2005 5–10 individuals or groups per km 50–99 Numerous, but < 5% cover or scat-

tered, with cover up to 5%3 Frequent 50

4 2–5 individuals or groups per km 20–49 Few, with small cover 2 Frequent 503 ± 1 individual or group per km 5–19 Solitary, with small cover 1 Occasional 102 Less abundant than above but more than 1

individual or group per 5 km2–4 Occasional 10

1 ± 1 plant or group per 5–10 km 1 Rare 1*, approximate numbers of individuals or groups per 10 km transect.†, very abundant extensive stands; abundant; many clumps or stands: frequent, many sightings of single plants or small groups: occasional, a few

sightings of one or a few plants: rare, one sighting of one or a few plants.#, weighted abundance, numbers of individuals or groups per 10 km transect/recording.

= +

Bothalia 37,2 (2007) 217

the results presented are not a true refl ection of their sta-tus (see asterisked species in Appendix 4). A further 45 species were recorded in the study area after 2000 and are asterisked in the species checklist (Appendix 5). A total of 601 species are listed in the full checklist given in Appendix 5—this is estimated to be about half the total number of naturalized and casual alien plant spe-cies in southern Africa. The most comprehensive listing of naturalized species in southern Africa, compiled by Wells et al. (1986), contains approximately 965 species, predominantly herbaceous. The SAPIA database, with a bias towards trees and shrubs, has an additional 231 spe-cies not listed by Wells et al. (1986).

Geographical extent of invasion

Alien plant invasion was recorded almost through-out the study area. Figure 1A shows invasion in terms of species numbers per QDS and Figure 1B shows the severity of invasion per QDS based on the total weighted abundance of all species per QDS. Most inva-sion, in terms of both species numbers and total species abundance, was recorded along the southern, southwest-ern and eastern coastal belts and in the adjacent interior. This corresponds with the regions of highest rainfall (Schulze 1997), urban development, and cultivation of agricultural and silvicultural crops. It also includes the whole of the Fynbos and Forest, and the moister east-ern parts of the Grassland and Savanna Biomes (Figure 1C). Distribution maps of 234 species, which include all declared species under the Conservation of Agricultural Resources Act, Act 43 of 1983, and amended in 2001, are given in the fi eld guide Alien weeds and invasive plants (Henderson 2001).

Prominent invasive species

There were 97 prominent invasive species in the study area and each of the biomes (Appendices 1–3). All these species were invading natural and semi-natu-ral habitats.

Study area

Fifty species account for most invasion (the upper 80% of the summed prominence values) in the study area (Appendix 1). Acacia mearnsii (black wattle) was the most prominent species by far, with a value of 18.37 (out of a maximum of 200) which is more than double the value of the second-ranked species, A. saligna (Port Jackson). The remaining top ten most prominent invad-ers in the study area were in order, Lantana camara (lantana), A. cyclops (rooikrans), Opuntia fi cus-indica (sweet prickly-pear), Solanum mauritianum (bugweed), Populus alba/×canescens (white/grey poplars—values of these two taxa were combined where they were diffi -cult to distinguish at a distance during roadside surveys), Melia azedarach (seringa), A. dealbata (silver wattle) and Prosopis spp. (P. glandulosa var. torreyana, P. velu-tina and their hybrids)(mesquite trees). Together these species cover almost the entire study area (Figures 2, 3).

Savanna Biome

Forty-eight species were the most prominent invaders in the Savanna Biome (Appendix 2). Lantana camara

was the most prominent species with a prominence value of 20.6, followed by Chromolaena odorata (triffi d weed) with a value of 14.2 and Melia azedarach with a value of 12. The remaining top ten invaders were, in order, Solanum mauritianum, Acacia mearnsii, Opuntia fi cus-indica, Ricinus communis (castor-oil plant), Psidium

FIGURE 1.—A, species numbers per quarter-degree square in study area; B, severity of invasion per quarter-degree square. Light invasion: < 1 individual or group per km. Moderate invasion: up to 5 individuals or groups per km; some species forming stands. Heavy invasion: up to 50 individuals or groups per km; many species forming stands; some completely dominating landscape. C, heavy invasion in relation to biomes in study area.

A

B

C

218 Bothalia 37,2 (2007)

guajava (guava), Eichhornia crassipes (water hyacinth) and Jacaranda mimosifolia (jacaranda).

Fynbos Biome

Twenty species were the most prominent invaders in the Fynbos Biome (Appendix 2). Acacia mearnsii was the most prominent species with a prominence value of 31.5, followed by A. saligna and A. cyclops with val-ues of 30.4 and 27.2, respectively. The remaining top ten most prominent invaders in order, were, Pinus pin-aster (cluster pine), Acacia melanoxylon (Australian

blackwood), A. longifolia (long-leaved wattle), Populus ×canescens (grey poplar), Paraserianthes lophantha (stinkbean), Rubus fruticosus (European blackberry) and Opuntia fi cus-indica. Hakea sericea (silky hakea) and Pinus radiata (radiata pine), both invaders of mountain fynbos, were most likely under-recorded because of the inacessibility and under-sampling of this habitat.

Forest habitats

Forty species were the most prominent invaders in forest habitats (Appendix 2). Chromolaena odorata was

FIGURE 2.—Distribution and severity of invasion in study area: A, Acacia cyclops; B, Acacia dealbata; C, Acacia mearnsii; D, Acacia saligna; E, Lantana camara; F, Melia azedarach. Light invasion, ; moderate invasion, ; heavy invasion, .

A B

C D

E F

Bothalia 37,2 (2007) 219

the most prominent species with a prominence value of 23.9, followed by Solanum mauritianum and Acacia mearnsii with values of 19 and 16.7, respectively. The remaining top ten prominent invaders were, in order, Acacia melanoxylon, Lantana camara, Cestrum laevi-gatum (inkberry), Caesalpinia decapetala (Mauritius/Mysore thorn), Melia azedarach, Pinus pinaster and Psidium guajava. Pereskia aculeata (pereskia) ranked eleventh and could have been vastly underestimated because of the diffi culty of observing this forest canopy, climbing species.

Grassland Biome

Thirty-two species were the most prominent invaders in the Grassland Biome (Appendix 3). Acacia mearnsii was the most prominent species with a prominence value of 21.3, followed by A. dealbata and Salix babylonica (weeping willow) with values of 20.9 and 17.3, respec-tively. The remaining top ten most prominent invaders were, in order, Populus alba/×canescens (white/grey poplars), Solanum mauritianum, Rubus spp. (mainly R. cuneifolius)(brambles), Pyracantha angustifolia and P. crenulata (yellow and Himalayan fi re thorns), Eucalyptus spp. (eucalypts), Melia azedarach and Opuntia fi cus-indica. Campuloclinium macrocephalum (pompom weed) which did not feature as a prominent invader in this study showed an explosive rate of increase after

2000 and currently would be rated as one of the most prominent invaders in the Grassland Biome (Henderson et al. 2003).

Nama-Karoo Biome

Fourteen species were the most prominent invaders in the Nama-Karoo Biome (Appendix 3). Prosopis spp. (P. glandulosa var. torreyana, P. velutina and their hybrids) were the most prominent species with a prominence value of 60.6, followed by Atriplex infl ata (sponge-fruit saltbush) and Opuntia fi cus-indica with values of 21 and 14 respectively. The remaining top ten prominent invad-ers were, in order, Salsola kali/tragus (Russian tumble-weed), Azolla fi liculoides (red water fern), Nicotiana glauca (wild tobacco), Atriplex nummularia (old man saltbush), Schinus molle (pepper tree), Agave americana (American agave) and Solanum elaeagnifolium (silver-leaf bitter-apple).

Succulent Karoo Biome

Twelve species were the most prominent invaders in the Succulent Karoo Biome (Appendix 3). Nicotiana glauca was the most prominent invader with a promi-nence value of 26.8, followed by Acacia cyclops and Prosopis spp. (P. glandulosa var. torreyana, P. velutina and their hybrids) with values of 26.3 and 25.9, respec-

FIGURE 3.—Distribution and severity of invasion in study area: A, Opuntia fi cus-indica; B, Populus alba/×canescens; C, Prosopis spp.; D, Solanum mauritianum. Light invasion, ; moderate invasion, ; heavy invasion, .

A B

C D

220 Bothalia 37,2 (2007)

tively. The remaining top ten most prominent invaders were, in order, Acacia mearnsii, A. saligna, Atriplex infl ata, Arundo donax (giant reed), Atriplex nummularia, Opuntia fi cus-indica and Populus ×canescens.

Riparian and wetland habitats

Fifty-fi ve species had more than 50 records in ripar-ian and wetland habitats (Appendix 4). Salix babylonica was the most frequently recorded riparian and wetland species with 1 323 records, followed by Populus alba/×canescens with 1 176 records and Acacia mearnsii with 953 records. The remaining top ten riparian and wetland invaders were, in order, Melia azedarach, Ricinus com-munis, Arundo donax, Acacia dealbata, Sesbania puni-cea (red sesbania), Prosopis spp. and Nicotiana glauca.

Biome comparison

The Savanna Biome, which occupies the largest number of QDS (645) in the study area, had the great-est number of species (358) and the most invasion in terms of total abundance of all species (Table 2). The Fynbos Biome, however, which occupies the least QDS (139), was the most heavily invaded in terms of average abundance of all species per QDS, average abundance of individual species per QDS and % QDS heavily invaded. The Grassland Biome ranks third after Fynbos for total abundance of all species, followed by Forest, Nama-Karoo and the Succulent Karoo Biome was the least invaded.

Biome profi les

Appendix 6 provides species characteristics of the prominent invasive species. Table 3 analyses the promi-

nent invasive species in each of the biomes and the study area in terms of region of origin, taxonomy, growth form, perennation, type of reproduction, dispersal mecha-nism and cultivated use.

Savanna Biome species are predominantly of tropical origin; members of the Fabaceae, Solanaceae, Asteraceae and Rosaceae; woody trees and shrubs, followed by herbs and climbers; perennial evergreen and evergreen/deciduous; seed-producers; water and bird dispersed; ornamentals and agricultural crops.

Fynbos Biome species are predominantly of temperate origin (particularly southern temperate); members of the Fabaceae, Myrtaceae, Pinaceae and Salicaceae; woody trees and shrubs; perennial evergreen; seed-producers; water, bird and wind dispersed; silvicultural crops, orna-mentals and cover/binders.

Forest habitat species are predominantly of tropical origin; members of the Fabaceae, Asteraceae, Myrtaceae, Solanaceae, Pinaceae and Zingiberaceae; woody trees and shrubs, followed by herbs and climbers; perennial evergreen; seed-producers; bird and water dispersed; ornamentals, barriers and silvicultural crops.

Grassland Biome species are predominantly of north-ern temperate origin and the tropics; members of the Rosaceae, Fabaceae and Salicaceae; woody trees and shrubs, followed by herbs; perennial evergreen/decidu-ous and deciduous; seed-producers, but a greater per-centage of species coppice and sucker than in other veg-etation categories; water and bird dispersed; barriers, ornamentals and agricultural crops.

Nama-Karoo Biome species are predominantly of northern temperate origin and the tropics; members of the Chenopodiaceae, Salicaceae, Cactaceae, Fabaceae, Solanaceae and Tamaricaceae; woody trees and shrubs, followed by herbs and succulent trees and shrubs; peren-nial evergreen/deciduous and deciduous; seed-producers, but a greater percentage of species reproduce by vegeta-tive division than in other vegetation categories; water and wind dispersed; agricultural crops and ornamentals.

Succulent Karoo Biome species are predominantly of temperate origin; members of the Fabaceae, Chenopo-diaceae and Tamaricaceae; woody trees and shrubs; perennial evergreen and evergreen/deciduous; seed-pro-ducers and reproduce vegetatively by coppicing; water and wind dispersed; agricultural crops, ornamentals and cover/binders.

DISCUSSION

Biome comparison: extent of invasion

No previous studies have enabled a direct comparison of the extent of invasion in the different biomes using the same parameters. This study reinforces previous studies that the Fynbos Biome is the most extensively invaded vegetation type in South Africa (Richardson et al. 1997) but it also shows that parts of Savanna and Grassland are

TABLE 2.—Biome comparison in terms of extent, numbers and abun-dance of species and severity of invasion

FB Fh SB GB NKB SKBExtent in QDS 139 157 645 521 548 141Total species 216 172 358 319 105 69Prominent invasive

species20 40 48 32 14 12

Total abundance* 986 653

222 419

1165 895

811 723

211 589

67 524

Ave abundance per QDS

7 098 1 417 1 808 1 558 386 479

Ave abundance per species

4 568 1 293 3 257 2 545 2 015 979

% QDS light# 17 15 51 49 51 66% QDS moderate# 30 44 20 33 13 9% QDS heavy# 47 41 8 9 1 1QDS, quarter-degree squares in Fynbos, Savanna, Grassland, Nama-

Karoo and Succulent Karoo according to Rutherford (1997); QDS in forest habitats according to SAPIA database.

*, total weighted abundance of all species (see text).Prominent invasive species: species with highest prominence values

adding up to ± upper 80% of summed values (see text).#, % QDS lightly invaded: less than 1 individual or group per km; #,

% QDS moderately invaded: up to 5 individuals or groups per km; some species forming stands; #, % QDS heavily invaded: up to 50 individuals or groups per km; many species forming stands; some completely dominating landscape.

FB, Fynbos Biome; Fh, Forest habitats; SB, Savanna Biome; GB, Grassland Biome; NKB, Nama-Karoo Biome; SKB, Succulent Karoo Biome.

Bothalia 37,2 (2007) 221

TABLE 3.—Analysis of region of origin, taxonomy, growth forms, perennation, reproduction, dispersal mechanisms and cultivated uses of promi-nent invasive species in each of the biomes, forest habitats and study area

Characteristics Savanna Biome (49 spp.)

Fynbos Biome (24 spp.)

Forest habitats (53 spp.)

Grassland Biome (35 spp.)

Nama-Karoo Biome (18 spp.)

Succulent Karoo Biome (16 spp.)

Study area (97 spp.)

Region of originNorthern temperate

spp. 10 (20%) 7 (29%) 14 (26%) 18 (51%) 9 (50%) 6 (38%) 32 (33%)

Southern temperate spp.

4 (8%) 10 (42%) 6 (11%) 3 (9%) 3 (17%) 5 (31%) 15 (15%)

Tropical spp. 34 (69%) 7 (29%) 33 (62%) 13 (37%) 6 (33%) 5 (31%) 49 (51%)Hybrid 1 (2%) 1 (3%) 1 (1%)TaxonomyFamilies 18 12 24 14 10 10 32Families with 50% or

more of total species (no. spp. in brackets)

Fabaceae (9); Solanaceae (7); Asteraceae (5); Rosaceae (4)

Fabaceae (8); Myrtaceae (3); Pinaceae (2); Salicaceae (2)

Fabaceae (7); Asteraceae (5); Myrtaceae (4); Solanaceae (4); Pinaceae (3); Zingiberaceae (3)

Rosaceae (9); Fabaceae (6) Salicaceae (6)

Chenopodiaceae (3); Salicaceae (3); Cactaceae (2); Fabaceae (2); Solanaceae (2); Tamaricaceae (2)

Fabaceae (5); Chenopodiaceae (2); Tamaricaceae (2)

Fabaceae (15); Rosaceae (9); Solanaceae (9); Asteraceae (6); Salicaceae (6); Cactaceae (5); Myrtaceae (5)

Growth form (spp.)Woody tree & shrub 26 (53%) 21 (88%) 33 (62%) 26 (74%) 9 (50%) 12 (75%) 60 (62%)Succulent tree &

shrub 5 (10%) 1 (4%) 1 (2%) 1 (3%) 3 (17%) 1 (6%) 6 (6%)

Climber 7 (14%) 0 8 (15%) 1 (3%) 0 0 9 (9%)Herbaceous 10 (20%) 1 (4%) 10 (19%) 6 (17%) 5 (28%) 2 (13%) 20 (21%)Grass/reed 1 (2%) 1 (4%) 1 (2%) 1 (3%) 1 (6%) 1 (6%) 2 (2%)Perennation (spp.)Perennial evergreen 20 (41%) 18 (75%) 33 (62%) 13 (37%) 6 (33%) 8 (50%) 51 (53%)Perennial evergreen/

deciduous 11 (22%) 3 (13%) 9 (17%) 8 (23%) 5 (28%) 5 (31%) 19 (20%)

Perennial deciduous 7 (14%) 2 (8%) 5 (9%) 9 (26%) 5 (28%) 2 (13%) 14 (14%)Variable 3 (6%) 1 (4%) 2 (4%) 2 (6%) 1 (6%) 1 (6%) 3 (3%)Germinative (annual/

biennial) 8 (16%) 0 4 (8%) 3 (9%) 1 (6%) 0 10 (10%)

Reproduction (spp.) by:Seeds/spores 44 (90%) 21 (88%) 50 (94%) 29 (83%) 14 (78%) 14 (88%) 91 (94%)Coppicing 23 (47%) 11 (46%) 25 (47%) 22 (63%) 8 (44%) 11 (69%) 44 (45%)Suckering 9 (18%) 3 (13%) 6 (11%) 9 (26%) 4 (22%) 2 (13%) 14 (14%)Division 6 (12%) 4 (17%) 3 (6%) 6 (17%) 5 (28%) 2 (13%) 10 (10%)Rhizomes 1 (2%) 1 (4%) 4 (8%) 1 (3%) 2 (11%) 1 (6%) 6 (6%)Stolons/runners 1 (2%) 0 3 (6%) 1 (3%) 0 0 3 (3%)Bulbils 1 (2%) 0 0 0 0 0 1 (1%)Dispersal (spp.) by: Wind 13 (27%) 7 (29%) 19 (36%) 4 (11%) 7 (39%) 6 (38%) 31 (32%)Water 27 (55%) 16 (66%) 22 (42%) 19 (54%) 10 (56%) 11 (69%) 47 (48%)Birds 17 (35%) 8 (33%) 25 (47%) 17 (49%) 5 (28%) 4 (25%) 40 (41%)Mammals 9 (18%) 5 (21%) 7 (13%) 3 (9%) 4 (22%) 5 (31%) 16 (16%)Humans 14 (29%) 5 (21%) 8 (15%) 10 (29%) 3 (17%) 3 (19%) 21 (22%)Ants 8 (16%) 3 (13%) 4 (8%) 6 (17%) 0 2 (13%) 10 (10%)Cultivated uses (spp.)Ornamental 20 (41%) 6 (25%) 21 (40%) 7 (20%) 5 (28%) 4 (25%) 35 (36%)Cover/binder 4 (8%) 5 (21%) 6 (11%) 4 (11%) 2 (11%) 3 (19%) 9 (9%)Barrier 5 (10%) 2 (8%) 9 (17%) 8 (23%) 2 (11%) 0 17 (18%)Silvicultural crop 2 (4%) 7 (29%) 8 (15%) 4 (11%) 0 2 (13%) 10 (10%)Agricultural crop 11 (22%) 4 (17%) 7 (13%) 7 (20%) 7 (39%) 7 (44%) 16 (16%)Species with no uses 7 (14%) 0 2 (4%) 5 (14%) 2 (11%) 0 11 (11%)

222 Bothalia 37,2 (2007)

as heavily invaded as parts of the Fynbos. These fi nd-ings have important implications for the management of alien plant invasions in South Africa. Without inter-vention we can expect invasion to increase in all parts of South Africa and particularly in the Grassland and Savanna Biomes where large areas are yet to be invaded and many species are only starting to invade.

Biome comparison: prominent invaders

Each biome has a different suite of prominent invad-ers. In part, this can be explained by their pre-adap-tion to the prevailing environmental conditions, but also to their history of planting. Most of these species were deliberately introduced and cultivated on a grand scale as silvicultural and agricultural crops e.g. Acacia mearnsii, A. melanoxylon, Pinus pinaster and species of Prosopis, as barriers e.g. Acacia dealbata, Hakea seri-cea and Pyracantha angustifolia, as cover/binders e.g. Acacia cyclops, A. saligna and Populus ×canescens, and ornamentals e.g. Melia azedarach and Lantana camara.

Some species which have become prominent invad-ers were not cultivated widely or on a grand scale e.g. Solanum mauritianum, Chromolaena odorata and Nicotiana glauca. Although the latter species have on occasion been cultivated as ornamentals they have man-aged to disperse very effi ciently without human assis-tance—C. odorata by wind, S. mauritianum by birds and N. glauca by wind, soil and water.

Some species, although widely planted, have become prominent invaders in only one biome, indicating that environmental factors have limited their distribution. Examples are members of the family Rosaceae, such as Pyracantha angustifolia, P. crenulata, Cotoneaster franchetii and C. pannosus that are virtually restricted to high-altitude grasslands where it appears that freez-ing winter temperatures are needed to trigger seed ger-mination (Henderson 1989). Jacaranda mimosifolia is another species that has been planted throughout South Africa yet is only invasive in the moister parts of the Savanna and Forest Biomes. In its native northeastern Argentina, J. mimosifolia occurs mainly on river banks under warmer-temperate, subhumid conditions (Poynton 1973)—environmental conditions which are similar to those in its naturalized range in southern Africa. A pre-vious study by Henderson (2006b) showed that the cur-rent distributions of invasive plants in southern Africa are a refl ection of the climatic zones of their origin.

There are considerable differences in the species pro-fi les of the biomes but shared features are the promi-nence of the family Fabaceae, woody trees and shrubs, reproduction by seed and water dispersal. Within the Fabaceae the Acacia species are the most numerous with 17 listed species and account for a very large pro-portion of all plant invasion in South Africa. They are important invaders of all the major vegetation types except for those in the arid interior, where other legu-minous invaders take over, namely species of Prosopis. The most widespread and abundant acacias are Acacia mearnsii, A. cyclops and A. saligna. Acacia mearnsii has invaded the widest range of vegetation types in South

Africa and is the most widespread riverine invader, occurring almost continuously from Louis Trichardt in the Limpopo Province down the eastern seaboard to Cape Town, a distance of ± 2 500 km. Acacia cyclops stretches along almost the entire Cape coastline from Port Nolloth in the northwest to beyond East London in the east, a distance exceeding 2 000 km. Acacia saligna stretches along the Cape coastline from Saldanha Bay in the west to the Kei River in the east, a distance of ± 1 500 km.

Sixty-eight per cent of prominent invaders are peren-nial trees or shrubs. There are only two grasses listed as prominent invaders and only 14 species as nonperennial (annual, biennial or variable). Grasses and herbaceous species are under-represented in the SAPIA database largely as a consequence of biassed recording of the larger, more conspicuous species. In southern Africa the Poaceae is one of the largest plant families with 847 indigenous species and 115 (12%) naturalized spe-cies (Gibbs Russell et al. 1990). However, only 30 grass species are listed in this publication. There is defi nitely a lack of expertise in identifying grasses in South Africa and this is one of the reasons for the under-represen-tation of alien grasses in weed surveys. There is simi-larly an under-representation of the alien herbaceous Asteraceae. The South African National Biodiversity Institute’s online species checklist at http://posa.sanbi.org/searchspp.php lists 125 alien herbaceous species in South Africa, yet only 44 alien herbaceous species have been listed in this publication.

Comparison with other studies

Versfeld et al. (1998) provide the only other assess-ment of the extent and importance of invasive plants on a national level. This study combined expert knowledge of local landowners and managers with existing data-bases such as those of provincial conservation authori-ties and national departments. The SAPIA database was used as a means of data verifi cation particularly for areas where expert knowledge was lacking. Overall the assessment by Versfeld et al. (1998) relating to impor-tance rankings and the distribution of dense infesta-tions concurs with this study. Eight of the top ten invad-ing species or groups of species, ranked by condensed invaded area, also appear within the top ten ranking in this study—these are: Acacia cyclops, Prosopis spp., A. mearnsii, A. saligna, Solanum mauritianum, Opuntia spp., Melia azedarach and Lantana camara. Versfeld et al. (1998) include Pinus spp. and Hakea spp. within the top ten ranking, whereas this study includes Populus alba/×canescens and Acacia dealbata. The lower rank-ing of Pinus spp. and Hakea spp. in this study can be explained by the under-sampling of mountain habitats, which are largely inaccessible by road, in which these species are invasive.

Abundance data presented in this study suggests that Versfeld et al. (1998) may have underestimated the area of invasion of Salix babylonica and Populus alba/×canescens. In the present study these species were not only the most frequently recorded invaders in riparian and wetland habitats but their total weighted abundance

Bothalia 37,2 (2007) 223

was in both instances more than Melia azedarch and Eucalyptus spp. (Appendix 1) which were rated above Salix spp. and Populus spp. by Versfeld et al. (1998). Other riparian species which may also have been under-estimated include Arundo donax (giant reed), Morus alba (common mulberry) and Ricinus communis.

Looking to the future

The Working for Water Programme (WfW) and biologi-cal control

Alien plant invasion is a dynamic process and there will undoubtedly be changes in species composi-tion and prominence of invaders in the future. Many of the large tree species—mainly Acacia, Eucalyptus, Pinus, Populus, Prosopis species and Melia azedarach have been targeted by a national clearing programme, Working for Water (WfW), which started in October 1995 (Marais et al. 2004). To date there has not been an assessment of the affects of the WfW programme on the status of invasive alien infestations. The programme has been proposed for 20 years but Marais et al. (2004) indicate that even with the existing generous levels of funding, it is unlikely that the problem will be contained within the next half century.

Biological control of invasive plants using introduced insects and pathogens is the only sustainable, effec-tive and inexpensive solution to the most intractable of the invasive alien plant problems (Marais et al. 2004). When they are successful, the damage infl icted by bio-logical control agents causes a decline in population densities, distribution and, or, rates of spread of inva-sive plants, and reduces the costs of other management practices (Zimmermann et al. 2004). There have been some outstanding successes with biocontrol in South Africa, dating back to the early and mid-1900s with Opuntia monacantha (drooping prickly pear) and O. fi cus-indica, and in more recent years with several of the Acacia spp. (Zimmermann et al. 2004). Population monitoring of A. saligna in the Western Cape has shown marked decreases in population densities caused by the gall-forming rust fungus, Uromycladium tepperianum (Morris 1997; Wood & Morris 2007).

New invaders

Since 2000 a further 45 species have been added to the SAPIA database for the study area (Appendix 5). Another eight species, two of which are indigenous to South Africa, are naturalized in neighbouring Zimbabwe and Malawi (Appendix 5). All but three of the additional species have been listed as weeds in A global compen-dium of weeds (Randall 2002) and 28 species are envi-ronmental weeds elsewhere in the world and therefore have the potential to become invasive in South Africa. Fourteen of the new species are ‘noxious weeds’ or restricted in California, Florida, Hawaii, New Zealand and Australia—places with similar climates and with which South Africa has many invasive species in com-mon. We should be especially wary of these species which include some of the most damaging and costly invaders such as Hydrilla verticillata (hydrilla), a sub-

merged aquatic plant that has invaded much of the USA since the 1960s, and Chondrilla juncea (skeleton weed), a terrestrial herb that has become a major agricultural weed in the USA, Canada and Australia.

CONCLUSION

The main objective of this paper was to provide a historical overview of the extent and species composi-tion of alien plant invasion in southern Africa from 1979 until the end of 2000. This snapshot of invasion will pro-vide a yardstick by which we can measure our progress or failure in the management of invasive alien plants in southern Africa.

This publication will also contribute to the global knowledge of invasive alien plants. One of the most useful predictors of invasiveness is whether a species is invasive elsewhere in the world (Richardson et al. 2004a). The lists of prominent invaders and other natu-ralized species provided here will serve as a warning to neighbouring countries and to those as far afi eld as Australia, New Zealand and the USA of potentially inva-sive species in their regions.

The results presented here are but a summary of the more than 50 000 records of invasive alien plants in the SAPIA database. Much more can be gleaned from the SAPIA data. SAPIA has provided the raw data for analyses that have been used to prioritize invasive alien species for management (Robertson et al. 2003; Nel et al. 2004), to map the potential spread of invasive plants (Rouget et al. 2004), to look at broad-scale distribution patterns of invasive species (Richardson et al. 2004b), to correlate patterns of alien plant species richness with the environment and indigenous species richness (Richardson et al. 2005), to correlate patterns of invasion with interactions between environment, species traits and human uses (Thuiller et al. 2006) and to look at potential range and residence time (Wilson et al. 2007). SAPIA has also played a crucial role in providing information on invasive plants for the revision of the Conservation of Agricultural Resources Act, Act 43 of 1983, and the drafting of the National Environmental Management: Biodiversity Act, Act 10 of 2004.

Alien plant invasion is a dynamic process and there-fore it is essential that the SAPIA database be kept up-to-date with current information. From October 2006 a sec-ond phase of the SAPIA mapping project was launched and all the SAPIA data will be available online at the Weeds and Invasive Plants (WIP) website, www.agis.agric.za/wip (Henderson 2006a).

ACKNOWLEDGEMENTS

I thank all the people who have contributed in many different ways to the compilation of the SAPIA data-base and have made it one of the most comprehensive databases on invasive alien plants in southern Africa. Les Powrie and Mike Rutherford of the South African National Biodiversity Institute are thanked for the QDS

224 Bothalia 37,2 (2007)

coverage of the biomes of South Africa which was used to subdivide the SAPIA dataset into the six biome data-sets for this publication. Special tribute is made to Mike Wells of the Botanical Research Institute for his inspi-ration and mentorship during the development of road-side survey techniques from 1979 to the mid-1980s, and to Helmuth Zimmermann of the Plant Protection Research Institute for his motivation and support of the SAPIA mapping project. SAPIA is an initiative of the Agricultural Research Council (ARC): Plant Protection Research Institute which has provided the infrastruc-ture, basal funding and support since the mid 1980s to the present. External funding of SAPIA has been grate-fully received from the Departments of Agriculture, Environmental Affairs and Tourism, and Water Affairs and Forestry (Working for Water Programme).

REFERENCES

GIBBS RUSSELL, G.E., WATSON, L., KOEKEMOER, M., SMOOK, L., BARKER, N.P., ANDERSON, H.M. & DALLWITZ, M.J. 1990. Grasses of southern Africa. Memoirs of the Botanical Survey of South Africa No. 58.

HENDERSON, L. 1989. Invasive alien woody plants of Natal and the north-eastern Orange Free State. Bothalia 19: 237–261.

HENDERSON, L. 1991a. Invasive alien plants of the Orange Free State. Bothalia 21: 73–89.

HENDERSON, L. 1991b. Invasive alien woody plants of the northern Cape. Bothalia 21: 177–189.

HENDERSON, L. 1992. Invasive alien woody plants of the eastern Cape. Bothalia 22: 119–143.

HENDERSON, L. 1995. Plant invaders of southern Africa. Plant Protection Research Institute Handbook No. 5. Agricultural Research Council, Pretoria.

HENDERSON, L. 1998a. Invasive alien woody plants of the south-ern and southwestern Cape region, South Africa. Bothalia 28: 91–112.

HENDERSON, L. 1998b. Southern African Plant Invaders Atlas (SAPIA). Applied Plant Sciences 12: 31, 32.

HENDERSON, L. 2001. Alien weeds and invasive plants. Plant Protection Research Institute Handbook No. 12. Agricultural Research Council, Pretoria.

HENDERSON, L. 2006a. Plant Protection Research Institute initia-tives: Southern African Plant Invaders Atlas (SAPIA) phase II. Plant Protection News 68: 5.

HENDERSON, L. 2006b. Comparisons of invasive plants in southern Africa originating from southern temperate, northern temperate and tropical regions. Bothalia 36: 201–222.

HENDERSON, L., GOODALL, J.M. & KLEIN, H. 2003. Pompom weed—an invader of grasslands that threatens conservation and agriculture in South Africa. Pamphlet produced by Gauteng Department of Agriculture, Conservation, Environment and Land Affairs (DACEL) in collaboration with the Agricultural Research Council (ARC).

HENDERSON, L. & MUSIL, K.J. 1984. Exotic woody plant invaders of the Transvaal. Bothalia 15: 297–313.

MARAIS, C., VAN WILGEN, B.W. & STEVENS, D. 2004. The clear-ing of invasive alien plants in South Africa: a preliminary assess-ment of costs and progress. South African Journal of Science 100: 97–103.

MORRIS, M.J. 1997. Impact of the gall-forming rust fungus Uromycladium tepperianum on the invasive tree Acacia saligna in South Africa. Biological Control 10: 75–82.

NEL, J.L., RICHARDSON, D.M., ROUGET, M., MGIDI, T.N., MDZEKE, N., LE MAITRE, D.C., VAN WILGEN, B.W., SCHONEGEVEL, L., HENDERSON, L. & NESER, S. 2004. A proposed classifi cation of alien invasive plant species in South Africa: towards prioritizing species and areas for management action. South African Journal of Science 100: 53–64.

POYNTON, R.J. 1973. Trees in South Africa—two hundred selected indigenous and exotic species: how to recognise and grow them. In W.F.E. Immelman, C.L. Wicht & D.P. Ackerman, Our green

heritage. Tafelberg, Cape Town.PYŠEK, P., RICHARDSON, D.M., REJMÁNEK, M., GRADY, L.,

WILLIAMSON, M. & KIRSCHNER, J. 2004. Alien plants in checklists and fl oras: towards better communication between taxonomists and ecologists. Taxon 53: 131–143.

RANDALL, R.P. 2002. A global compendium of weeds (Draft). Hawaiian ecosystems at risk. Available at www.hear.org/gcw.

RICHARDSON, D.M., MACDONALD, I.A.W., HOFFMANN, J.H. & HENDERSON, L. 1997. Alien plant invasions. In R.M. Cowling, D.M. Richardson & S.M. Pierce, Vegetation of southern Africa. Cambridge University Press, Cambridge.

RICHARDSON, D.M., PYŠEK, P., REJMÁNEK, M., BARBOUR, M.G., PANETTA, F.D. & WEST, C.J. 2000. Naturalization and invasion of alien plants: concepts and defi nitions. Diversity and Distributions 6: 93–107.

RICHARDSON, D.M., MORAN, V.C., LE MAITRE, D.C., ROUGET, M. & FOXCROFT, L.C. 2004a. Recent developments in the science and management of invasive alien plants: connecting the dots of research knowledge, and linking disciplinary boxes. South African Journal of Science 100: 126–128.

RICHARDSON, D.M., ROUGET, M., HENDERSON, L. & NEL, J.L. 2004b. Invasive alien plants in South Africa: macroecological patterns, with special emphasis on the Cape Floristic region. In M. Arianoutsou & V. Papanastasis, Proceedings of the 10th MEDECOS Conference, April 25–May 1, 2004, Rhodes, Greece. Millpress, Rotterdam.

RICHARDSON, D.M., ROUGET, M., RALSTON, S.J., COWLING, R.M., VAN RENSBURG, B.J. & THUILLER, W. 2005. Species richness of alien plants in South Africa: environmental corre-lates and the relationship with indigenous plant species richness. Ecoscience 12: 391–402.

ROBERTSON, M.P., VILLET, M.H., FAIRBANKS, D.H.K., HENDERSON, L., HIGGINS, S.I., HOFFMANN, J.H., LE MAITRE, D.C., PALMER, A.R., RIGGS, I., SHACKLETON, C.M. & ZIMMERMANN, H.G. 2003. A proposed prioritization system for the management of invasive alien plants in South Africa. South African Journal of Science 99: 37–43.

ROUGET, M., RICHARDSON, D.M., NEL, J.L., LE MAITRE, D.C., EGOH, B. & MGIDI, T. 2004. Mapping the potential spread of major plant invaders in South Africa using climatic suitability. Diversity and Distributions 10: 475–484.

RUTHERFORD, M.C. 1997. Categorization of biomes. In R.M. Cowling, D.M. Richardson & S.M. Pierce, Vegetation of southern Africa. Cambridge University Press, Cambridge.

SCHULZE, R.E. 1997. Climate. In R.M. Cowling, D.M. Richardson & S.M. Pierce, Vegetation of southern Africa. Cambridge University Press, Cambridge.

THUILLER, W., RICHARDSON, D.M., ROUGET, M., PROCHEŞ, Ş. & WILSON, J.R.U. 2006. Interactions between environment, species traits, and human uses describe patterns of plant inva-sions. Ecology 87: 1755–1769.

VERSFELD, D.B., LE MAITRE, D.C. & CHAPMAN, R.A. 1998. Alien invading plants and water resources in South Africa: a prelimi-nary assessment. WRC Report No. TT 99/98. Water Research Commission, Pretoria.

WELLS, M.J., BALSINHAS, A.A., JOFFE, H., ENGELBRECHT, V.M., HARDING, G. & STIRTON, C.H. 1986. A catalogue of problem plants in southern Africa. Memoirs of the Botanical Survey of South Africa No. 53.

WELLS, M.J., DUGGAN, K.J. & HENDERSON, L. 1980. Woody plant invaders of the central Transvaal. Proceedings of the third National Weeds Conference of South Africa, 1979. Balkema, Cape Town.

WILSON, J.R.U., RICHARDSON, D.M., ROUGET, M., PROCHEŞ, Ş., AMIS, M.A., HENDERSON, L. & THUILLER, W. 2007. Residence time and potential range: crucial considerations in modelling plant invasions. Diversity and Distributions 13: 11–22.

WOOD, A.R. & MORRIS, M.J. 2007. Impact of the gall-forming rust fungus Uromycladium tepperianum on the invasive tree Acacia saligna in South Africa: 15 years of monitoring. Biological Control 41: 68–77.

ZIMMERMANN, H.G., MORAN, V.C. & HOFFMANN, J.H. 2004. Biological control in the management of invasive alien plants in South Africa, and the role of the Working for Water Programme. South African Journal of Science 100: 34–40.

Bothalia 37,2 (2007) 225

Acaciacyclops 31 20 181 28 557 3.40 102 63 810 174 964 27.20 5 0 9 214 0.73dealbata 39 13 132 12 207 1.74 7 3 23 4 887 0.76 12 3 12 3 361 2.36decurrens 19 7 38 1 779 0.35 4 1 4 1 061 0.76longifolia 27 7 66 9 183 1.14 47 24 264 39 533 7.09 13 3 13 1 642 1.65mearnsii 134 59 710 75 210 10.20 88 70 871 210 388 31.50 36 22 91 22 825 16.70melanoxylon 33 2 73 1 274 0.49 50 23 298 38 206 7.35 25 12 85 18 186 14.20

Acaciacyclops 166 91 1 097 203 636 8.66 4dealbata 256 115 1 079 133 146 6.45 9decurrens 101 31 232 23 456 1.23longifolia 94 34 363 53 080 2.43mearnsii 428 251 2 620 410 950 18.37 1melanoxylon 134 29 482 43 926 2.40pycnantha 35 15 135 13 864 1.77saligna 158 93 1 030 219 223 9.00 2

Achyranthes aspera 77 3 85 1 262 0.22Agave

americana 431 8 761 9 136 1.94sisalana 170 12 293 8 924 0.91

Ageratumconyzoides 37 8 50 5 141 0.27conyzoides/houstonianum 31 8 49 2 994 0.20houstonianum 26 2 31 1 728 0.12

Argemonemexicana 27 5 36 2 134 0.14ochroleuca 154 20 206 12 115 0.82sp. 14 2 15 1 338 0.07

Arundo donax 371 82 855 50 158 3.41Atriplex

infl ata 164 77 213 58 878 2.28nummularia 172 18 333 10 248 1.04

Azolla fi liculoides 194 92 354 47 220 2.23Caesalpinia decapetala 127 41 413 33 868 1.94Cardiospermum

grandifl orum 43 12 57 4 905 0.28grandifl orum/halicacabum

16 5 20 2 427 0.12

Cereus jamacaru 124 11 193 13 042 0.82Cestrum laevigatum 70 16 167 11 039 0.70Chromolaena odorata 93 64 558 137 654 5.46Cinnamomum camphora 10 1 19 352 0.05Cirsium vulgare 188 20 345 14 022 1.18Datura

ferox 175 14 201 8 685 0.71innoxia 29 3 36 1 897 0.14sp. 84 1 110 1 710 0.29stramonium 286 24 373 12 932 1.21

Eichhornia crassipes 87 72 431 79 893 3.40Eucalyptus

camaldulensis 121 22 182 13 259 0.80diversicolor 49 8 153 5 562 0.50grandis 100 16 190 14 475 0.86sp. 505 30 1 103 23 523 3.12

Hakea sericea 77 17 230 15 959 0.99Ipomoea

indica 23 3 27 740 0.08indica/purpurea 74 7 120 2 284 0.33purpurea 37 3 46 1 801 0.16sp. 3 0 3 52 0.01

Jacaranda mimosifolia 195 16 613 17 430 1.87

Lantana camara 247 116 2 111 140 496 8.92 3Leptospermum laevigatum 38 15 102 15 916 0.71Litsea glutinosa 8 3 10 2 713 0.11Macfadyena unguis-cati 22 9 52 9 531 0.41Melia azedarach 551 65 2 119 65 735 6.64 8Morus alba 129 8 304 5 983 0.85Nephrolepis exaltata 13 1 19 459 0.06Nicotiana glauca 383 14 957 22 132 2.76Opuntia

fi cus-indica 861 57 2 445 72 477 7.55 5robusta 225 2 337 3 244 0.83stricta 106 14 193 5 412 0.59

Paraserianthes lophantha 54 9 286 20 042 1.24Passifl ora edulis 32 0 55 261 0.13Pennisetum clandestinum 48 12 53 8 884 0.39Pereskia aculeata 44 8 102 5 788 0.40Pinus

patula 85 13 238 11 636 0.88pinaster 85 44 401 48 229 2.36radiata 70 15 206 9 243 0.73

Populusalba 15 2 22 915 0.08 7alba/canescens 185 47 460 33 871 2.04 7×canescens 371 130 939 87 397 4.74 7

Prosopisglandulosa 40 10 50 4 988 0.26 10glandulosa/velutina 390 78 1 107 92 751 5.27 10velutina 48 6 53 3 108 0.21 10

Prunus persica 319 1 728 7 401 1.81Psidium guajava 160 50 732 55 791 3.31Pyracantha angustifolia 142 3 285 3 735 0.74Pyracantha angustifolia/cre-

nulata40 3 51 1 183 0.15

Ricinus communis 456 56 1 701 48 855 5.21Robinia pseudoacacia 110 14 178 8 828 0.66Rosa rubiginosa 119 12 276 11 494 0.95Rubus

cuneifolius 75 35 236 49 313 2.03fruticosus 89 32 244 22 810 1.23pascuus 3 2 3 450 0.02sp. 86 30 179 29 694 1.30×proteus 4 3 4 650 0.03

Salixbabylonica 475 89 1 381 85 116 5.63fragilis 75 24 176 15 710 0.87

Salsola kali/tragus 155 31 187 14 080 0.84Schinus molle 231 2 407 5 355 1.05Senna didymobotrya 139 29 339 16 342 1.24Sesbania punicea 323 68 830 52 078 3.41Solanum

elaeagnifolium 51 11 60 14 136 0.57mauritianum 265 99 1 364 135 219 7.14 6seaforthianum 30 3 77 2 656 0.25

Tithonia diversifolia 49 5 123 4 085 0.39Xanthium strumarium 149 21 212 12 633 0.85

APPENDIX 1.—Prominent invaders in study area

Scientifi c name QSp QSa Tr A Pv R Scientifi c name QSp QSa Tr A Pv R

APPENDIX 2.—Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats

Savanna Biome Fynbos Biome Forest HabitatsScientifi c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv

Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter-degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text).

Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation.QSp, quarter-degree squares present; QSa, quarter-degree squares abundant; Tr, total records; A, total weighted abundance (see text); Pv,

prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text); R, ranking of top ten taxa (taxa that are diffi cult to distinguish are grouped together).

226 Bothalia 37,2 (2007)

Acacia (cont.)pycnantha 3 1 4 203 0.04 32 14 131 13 661 2.92saligna 27 11 96 12 170 1.55 105 74 860 200 582 30.40 4 2 8 1 313 1.15

Achyranthes aspera 40 3 44 1 078 0.32 5 0 5 5 0.06 10 2 12 468 1.05Agave

americana 146 3 267 3 986 1.75 39 0 84 427 1.02 2 0 2 51 0.16sisalana 132 12 251 8 801 2.08 10 0 10 46 0.12 3 0 4 22 0.29

Ageratumconyzoides 32 8 44 4 979 0.66 1 0 1 1 0.07conyzoides/houstonianum 27 6 45 2 583 0.46 1 0 1 1 0.01 12 3 13 726 1.24houstonianum 22 2 26 1 625 0.28 8 2 8 1 261 1.13

Argemonemexicana 23 4 32 1 082 0.26 1 1 1 1 000 0.11 1 0 1 50 0.09ochroleuca 86 13 119 7 190 1.24 2 1 2 201 0.04 1 0 1 50 0.09sp. 6 2 7 1 272 0.15

Arundo donax 132 34 372 20 126 3.69 71 25 172 16 721 3.70 3 1 3 1 020 0.67Atriplex

infl ata 13 9 16 7 701 0.75 19 10 25 7 148 1.02nummularia 11 3 18 2 336 0.30 24 1 45 851 0.61

Azolla fi liculoides 43 16 75 4 979 0.83 16 10 37 7 491 1.19 2 2 2 400 0.32Caesalpinia decapetala 80 22 239 20 966 3.06 17 7 25 6 053 4.48Cardiospermum

grandifl orum 40 11 54 4 703 0.69 5 3 7 1 511 1.17grandifl orum/halicacabum 14 5 18 2 416 0.30 4 2 4 1 251 0.84

Cereus jamacaru 100 11 169 12 857 1.99 6 0 6 6 0.07 1 0 1 10 0.07Cestrum laevigatum 48 12 138 7 551 1.38 6 0 9 58 0.11 22 9 30 6 457 5.01Chromolaena odorata 79 57 529 133 524 14.20 33 25 77 41 029 23.90Cinnamomum camphora 7 1 13 346 0.10 2 0 3 3 0.04 7 1 14 347 1.14Cirsium vulgare 43 4 68 1 708 0.51 8 0 8 75 0.10Datura

ferox 69 3 87 2 676 0.69 3 0 3 3 0.04 2 0 2 11 0.15innoxia 23 1 30 1 435 0.28 1 0 1 50 0.02 1 0 1 50 0.09sp. 34 0 41 581 0.27 1 1 1 200 0.03stramonium 104 9 131 4 082 1.04 15 1 19 1 357 0.36 6 1 6 1 170 0.95

Eichhornia crassipes 46 44 279 60 302 6.64 20 7 46 5 591 1.10Eucalyptus

camaldulensis 23 4 36 1 993 0.36 41 13 81 6 318 1.59 3 2 3 2 050 1.13diversicolor 1 0 1 1 0.01 44 7 148 5 349 2.27 12 4 28 1 897 2.82grandis 54 8 111 9 827 1.43 1 0 1 10 0.01 15 2 19 974 1.77sp. 122 8 299 7 242 2.20 72 10 147 4 838 2.21 8 2 17 687 1.51

Hakea sericea 7 2 14 537 0.12 61 14 204 14 344 3.84 3 0 3 61 0.24Ipomoea

indica 18 3 22 735 0.18 3 0 3 3 0.04 3 0 3 3 0.21indica/purpurea 49 3 86 1 101 0.55 10 1 18 409 0.25 11 2 16 1 308 1.71purpurea 18 1 23 410 0.16 7 2 10 1 271 0.25 7 1 9 1 102 1.13sp. 2 0 2 51 0.01

Jacaranda mimosifolia 139 16 497 16 767 4.06 17 2 21 1 589 2.19Lantana camara 162 90 1 843 126 418 20.60 25 2 60 3 462 1.05 40 28 63 17 136 12.10Leptospermum laevigatum 3 3 6 701 0.09 35 12 96 15 215 2.66 2 0 3 21 0.22Litsea glutinosa 8 3 10 2 713 0.29 4 2 6 2 451 1.52Macfadyena unguis-cati 17 6 47 8 880 1.01 6 5 13 4 912 3.12Melia azedarach 291 53 1 394 54 100 12.00 44 0 82 181 0.98 17 5 29 4 856 4.22Morus alba 72 6 192 3 986 1.35 2 1 2 11 0.02 6 1 10 481 0.92Nephrolepis exaltata 10 1 12 318 0.09 2 0 6 131 0.08 9 1 13 337 1.07Nicotiana glauca 126 7 274 7 812 2.11 51 2 168 3 657 2.33 1 0 1 50 0.09Opuntia

fi cus-indica 330 39 1 159 47 136 10.10 73 5 267 7 242 3.85 8 5 8 4 261 2.48robusta 50 0 61 191 0.34 12 1 13 1 048 0.26stricta 82 13 168 5 112 1.32 1 0 1 50 0.02

Paraserianthes lophantha 5 0 10 104 0.06 47 9 274 19 936 5.22 7 0 7 34 0.51Passifl ora edulis 22 0 40 192 0.23 5 0 6 24 0.07 13 0 19 64 1.37Pennisetum clandestinum 10 3 12 2 414 0.27 26 4 28 2 106 0.54 3 3 3 3 000 1.56Pereskia aculeata 34 7 91 5 569 0.96 4 0 4 13 0.05 14 4 39 2 963 4.08Pinus

patula 30 7 90 6 154 1.00 14 5 18 1 194 1.80pinaster 13 5 34 6 549 0.74 66 36 355 39 368 8.14 9 4 29 4 586 4.10radiata 3 0 4 53 0.03 53 15 186 8 902 3.08 4 3 20 1 235 1.96

Populusalba 6 0 11 350 0.09 1 0 1 50 0.02 alba/canescens 51 22 171 15 598 2.24 1 0 1 50 0.02 8 2 8 611 0.84×canescens 36 8 76 5 390 0.86 82 34 279 24 456 5.74 3 0 4 22 0.29

Prosopisglandulosa 7 5 13 1 421 0.19 2 0 2 51 0.03

APPENDIX 2.—Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats (cont.)



Bothalia 37,2 (2007) 227

Prosopis (cont.)glandulosa/velutina 64 5 168 4 869 1.30 22 2 63 1 657 0.90velutina 6 1 7 257 0.06 3 0 3 52 0.04

Prunus persica 53 0 115 933 0.69 41 0 65 191 0.78 1 0 1 1 0.07Psidium guajava 124 45 662 53 388 8.07 6 0 9 45 0.11 14 8 25 5 174 4.09Pyracantha

angustifolia 6 0 13 40 0.07 5 0 6 15 0.07 2 0 2 2 0.14angustifolia/crenulata 5 0 5 5 0.03 2 0 2 2 0.02 1 0 1 50 0.09

Ricinus communis 256 46 1 230 40 996 10.00 87 7 250 4 189 3.35 25 5 30 1 618 2.84Robinia pseudoacacia 10 0 13 80 0.08 6 0 6 15 0.07 1 0 1 1 0.07Rosa rubiginosa 8 0 12 128 0.07 4 0 6 162 0.09 2 0 2 51 0.16Rubus

cuneifolius 18 4 45 10 711 1.16 5 3 6 2 451 1.52fruticosus 15 4 24 2 146 0.31 55 21 188 16 874 3.91 7 3 14 1 701 1.75pascuus 2 1 2 250 0.03 sp. 31 7 88 8 076 1.16 5 2 5 430 0.10 10 1 12 443 1.04×proteus 2 2 2 400 0.05

Salixbabylonica 67 5 140 3 609 1.05 38 3 74 1 838 1.05 6 0 6 211 0.52fragilis 2 0 5 121 0.04 1 1 1 200 0.03

Salsola kali/tragus 22 2 27 1 027 0.23 12 1 13 511 0.20Schinus molle 52 0 82 463 0.47 24 0 49 358 0.61 1 0 1 10 0.07Senna didymobotrya 103 25 261 14 239 2.60 3 0 3 3 0.04 12 1 15 406 1.24Sesbania punicea 139 26 405 21 438 3.97 60 19 175 17 026 3.77 7 0 8 66 0.59Solanum

elaeagnifolium 18 0 18 183 0.11 3 1 3 251 0.06mauritianum 123 52 748 77 619 10.60 28 2 86 2 035 1.21 66 32 97 27 090 19.00seaforthianum 28 3 75 2 636 0.62 1 0 1 10 0.01 9 0 31 263 2.30

Tithonia diversifolia 46 5 120 4 033 0.98 8 0 10 313 0.84Xanthium strumarium 74 16 126 8 577 1.40 2 1 3 251 0.06 4 0 4 62 0.31Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter-

degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text).

APPENDIX 2.—Prominent invaders in Savanna Biome, Fynbos Biome and Forest habitats (cont.)


Acaciacyclops 1 0 1 10 0.04 32 8 105 12 150 26.30dealbata 206 99 922 116 050 20.90 2 0 2 2 0.06decurrens 82 24 194 21 677 4.06longifolia 19 3 31 4 362 0.76 1 0 2 2 0.16mearnsii 197 116 1 003 114 979 21.30 9 6 36 10 373 18.20melanoxylon 49 4 109 4 435 1.33 2 0 2 11 0.18saligna 2 0 2 11 0.02 24 8 72 6 471 15.30

Achyranthes aspera 30 0 34 128 0.26 2 0 2 51 0.09Agave

americana 148 2 219 2 507 1.87 81 3 160 1 961 5.90 17 0 31 255 2.84sisalana 21 0 25 52 0.19 3 0 3 21 0.10 4 0 4 4 0.32

Ageratumconyzoides 5 0 6 162 0.06conyzoides/houstonianum 3 2 3 410 0.07houstonianum 4 0 5 103 0.05

Argemonemexicana 3 0 3 52 0.03ochroleuca 38 2 56 1 155 0.54 26 4 27 3 549 2.52 2 0 2 20 0.19sp. 1 0 1 10 0.01 1 0 1 1 0.03 6 0 6 55 0.56

Arundo donax 89 8 188 5 068 1.97 51 8 77 2 927 3.80 28 7 46 5 316 11.50Atriplex

infl ata 3 1 4 460 0.09 83 42 107 37 882 21.00 46 15 61 5 687 13.30nummularia 4 0 6 153 0.06 88 11 181 4 732 7.90 45 3 83 2 176 9.81

Azolla fi liculoides 88 41 172 21 806 3.92 47 25 70 12 944 8.30Caesalpinia decapetala 47 19 174 12 902 2.83Cardiospermum

grandifl orum 3 1 3 202 0.05grandifl orum/halicacabum 2 0 2 11 0.02

Cereus jamacaru 12 0 12 106 0.10 6 0 6 73 0.22Cestrum laevigatum 16 4 20 3 430 0.57

APPENDIX 3.—Prominent invaders in Grassland Biome, Nama-Karoo Biome and Succulent Karoo Biome

Grassland Biome Nama-Karoo Biome Succulent Karoo Biome

Scientifi c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv


228 Bothalia 37,2 (2007)

Chromolaena odorata 14 7 29 4 130 0.72Cinnamomum camphora 1 0 3 3 0.02Cirsium vulgare 126 14 253 10 776 3.14 11 2 16 1 463 1.19Datura

ferox 82 9 85 3 429 1.03 21 2 26 2 566 2.03innoxia 3 2 3 401 0.68 2 0 2 11 0.07sp. 46 0 65 877 0.57 2 0 2 2 0.06 1 0 1 50 0.15stramonium 126 12 174 6 224 2.01 40 2 48 1 259 2.10 1 0 1 10 0.09

Eichhornia crassipes 21 21 106 14 000 2.48Eucalyptus

camaldulensis 16 3 17 2 406 0.42 14 2 14 2 195 1.48 27 0 34 347 3.21diversicolor 2 1 2 210 0.04 2 0 2 2 0.16grandis 45 8 78 4 638 1.13sp. 276 12 614 11 153 5.76 21 0 27 188 0.93 14 0 16 102 1.42

Hakea sericea 1 0 1 50 0.01 8 1 11 1 028 2.40Ipomoea

indica 2 0 2 2 0.01indica/purpurea 15 3 16 774 0.21purpurea 12 0 13 120 0.11sp. 1 0 1 1 0.01

Jacaranda mimosifolia 55 0 115 662 0.90 1 0 1 1 0.03Lantana camara 59 24 207 10 606 2.79 1 0 1 10 0.09Macfadyena unguis-cati 5 3 5 651 0.12Melia azedarach 177 12 588 11 198 5.58 36 0 49 250 1.65 3 0 6 6 0.49Morus alba 54 2 109 1 985 1.02 1 0 1 1 0.03Nephrolepis exaltata 1 0 1 10 0.01Nicotiana glauca 48 1 72 1 266 0.67 92 2 206 3 980 8.30 66 2 237 5 417 26.80Opuntia

fi cus-indica 257 10 570 11 437 5.48 161 3 368 5 688 14.00 40 0 81 974 7.87robusta 72 1 120 927 0.97 80 0 130 1 029 4.60 11 0 13 49 1.10stricta 12 1 12 229 0.11 9 0 9 18 0.29 2 0 3 3 0.24

Paraserianthes lophantha 2 0 2 2 0.16Passifl ora edulis 5 0 9 45 0.07Pennisetum clandestinum 10 4 11 3 354 0.49 2 1 2 1 010 0.54Pereskia aculeata 6 1 7 206 0.08 Pinus

patula 55 6 148 5 482 1.73pinaster 4 1 4 230 0.06 2 2 8 2 082 3.72radiata 8 0 10 86 0.08 6 0 6 202 0.78

Populusalba 8 2 10 515 0.13alba/canescens 128 25 283 18 169 4.26 5 0 5 54 0.18×canescens 198 75 486 51 371 9.80 40 8 67 4 537 4.20 15 5 31 1 643 4.89

Prosopisglandulosa 3 0 3 61 0.03 25 4 29 2 395 2.00 3 1 3 1 060 1.81glandulosa/velutina 29 6 54 4 738 0.97 214 55 666 73 664 56.00 61 10 156 7 823 24.00velutina 1 0 1 1 0.01 37 5 41 2 788 2.60 1 0 1 10 0.09

Prunus persica 211 1 530 6 232 4.56 13 0 15 33 0.49 1 0 3 12 0.26Psidium guajava 29 5 60 2 357 0.72 1 0 1 1 0.08Pyracantha

angustifolia 122 3 256 3 661 6.07 9 0 10 19 0.32angustifolia/crenulata 33 3 44 1 176 0.46

Ricinus communis 80 3 166 3 149 1.57 13 0 24 208 0.85 20 0 31 313 2.92Robinia pseudoacacia 83 14 145 8 576 2.09 11 0 14 157 0.51Rosa rubiginosa 104 12 255 11 201 3.20 3 0 3 3 0.10Rubus

cuneifolius 57 31 191 38 602 6.12fruticosus 15 5 24 3 219 0.57 4 2 8 571 1.48pascuus 1 1 1 200 0.03sp. 50 21 86 21 188 3.23×proteus 2 1 2 250 0.05

Salixbabylonica 310 79 1 069 78 092 17.30 54 2 90 1 555 3.55 6 0 8 26 0.67fragilis 71 23 169 15 388 3.10 1 0 1 1 0.03

Salsola kali/tragus 20 1 23 610 0.24 95 26 117 12 309 9.50 6 1 7 233 0.90Schinus molle 54 0 73 360 0.57 75 2 156 3 952 6.80 26 0 47 222 4.06Senna didymobotrya 33 4 75 2 100 0.79Sesbania punicea 117 22 238 13 269 3.34 4 0 4 13 0.13 3 1 8 332 1.13Solanum

elaeagnifolium 18 4 21 3 498 0.58 10 6 16 10 202 5.30 2 0 2 2 0.16mauritianum 114 45 530 55 565 10.60seaforthianum 1 0 1 10 0.01

Tithonia diversifolia 3 0 3 52 0.03Xanthium strumarium 64 2 72 2 364 0.81 9 2 11 1 441 1.03Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation. QSp, quarter-degree squares present; QSa, quarter-

degree squares abundant; R, records; A, total weighted abundance (see text); Pv, prominence value (bold numbers: highest prominence values which add up to ± upper 80% of summed values—see text).

APPENDIX 3.—Prominent invaders in Grassland Biome, Nama-Karoo Biome and Succulent Karoo Biome (cont.)

Grassland Biome Nama-Karoo Biome Succulent Karoo Biome

Scientifi c name QSp QSa R A Pv QSp QSa R A Pv QSp QSa R A Pv

Bothalia 37,2 (2007) 229

Acaciabaileyana 86 135 20 39 1 75 1 10cultriformis # 1 1 1cyclops 166 91 1 097 181 810 9 1 105 206dealbata 256 115 1 079 132 23 12 922 2 4 542decurrens 101 31 232 38 4 194 30elata 35 1 60 4 51 4 3 2 4fi mbriata 1 1 1 1implexa 2 2 2 2longifolia 94 34 363 66 264 13 31 2 132mearnsii 428 251 2 620 710 871 91 1 003 36 953melanoxylon 134 29 482 73 298 85 109 2 130paradoxa 1 2 2podalyriifolia 57 2 101 49 27 1 25 6pycnantha 35 15 135 4 131 6saligna 158 93 1 030 96 860 8 2 72 363viscidula 1 1 1

Acanthocereus ?tetragonus 1 1 1 1Acanthospermum

australe * 1 1 1hispidum * 1 1 1

Acanthus polystachyus var. pseudopubescens #

1 1 1

Acernegundo 1 1 1 1 1? sp. 1 1 1 1 1

Achyranthes aspera * 77 3 85 44 5 12 34 2 23Acorus calamus * 1 1 1 1Acrocarpus fraxinifolius # 1 1 1 1Adiantum raddianum # 1 1 1 1Agave

americanavar. americana 431 8 761 267 84 2 219 160 31 118var. expansa 1 1 1

decipiens # 1 1 1sisalana 170 12 293 251 10 4 25 3 4 13sp. 31 1 60 53 4 3 12

Ageratinaadenophora 11 4 26 11 10 3 5 10riparia ?# 1 1 1

Ageratumconyzoides 37 8 50 44 1 6 32conyzoides/houstonianum 31 8 49 45 1 13 3 24houstonianum 26 2 31 26 8 5 7

Agrimonia cf. parvifl ora # 1 1 1 1Agrostemma githago * 1 1 1Ailanthus altissima 32 2 40 11 6 19 3 1 9Albizia

chinensis # 1 1 1lebbeck 4 2 5 5procera 1 1 1

Alhagi maurorum 10 11 8 1 1 1 3Alisma plantago-aquatica 8 1 9 2 7 9Alnus glutinosa 1 1 1 1Alpinia zerumbet 5 7 2 4 1 1 2Alternanthera pungens * 4 4 1 3 1Amaranthus

hybridus * 3 3 3 1sp.* 1 1 1

Ambrosia artemisiifolia * 2 2 1 1 1Ammi majus * 1 1 1Anigozanthos fl avidus # 1 1 1 1Anredera cordifolia 24 2 25 17 3 7 5 4Antigonon leptopus 3 5 5 1Apium graveolens * 1 1 1

APPENDIX 4.—Summary of results for all naturalized and casual alien plants in the study area, Savanna Biome, Fynbos Biome, Forest habitats, Grassland Biome, Nama-Karoo Biome, Succulent Karoo Biome and watercourse/wetland habitats

Plant name QSp QSa Study area

records

Savanna Biome records

Fynbos Biome records

Forest habitat records

Grassland Biome records

Nama-Karoo Biome records

Succulent Karoo Biome records

Watercourse/wetland records

Combined taxa e.g. Ageratum conyzoides/houstonianum indicate uncertainty of identifi cation.#, casual alien plants: occurring outside cultivation; some species fl ourishing but less than 10 years of records in SAPIA precludes being categorized

as ‘naturalized’ (Pyšek et al. 2004).*, mainly herbaceous species that are suspected of being under-estimated in this survey.QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant.Bold numbers in biome categories add up to upper 80% or more of total records.

230 Bothalia 37,2 (2007)

Araujia sericifera 36 1 53 15 2 4 35 1 12Ardisia crenata 2 1 3 3 2 3Argemone

mexicana 27 5 36 32 1 1 3 20ochroleuca subsp. ochroleuca 154 20 206 119 2 1 56 27 2 50sp. 14 2 15 7 1 1 6 9

Aristolochia elegans 6 1 8 7 2 1 4Arundo donax 371 82 855 372 172 3 188 77 46 548Astartea fascicularis # 1 1 1Atriplex

infl ata 164 77 213 16 25 4 107 61 16muelleri * 1 1 1nummularia subsp. nummularia 172 18 333 18 45 6 181 83 107semibaccata * 4 5 3 2 1sp. 10 10 8 2 2

Azollafi liculoides 194 92 354 75 37 2 172 70 354?pinnata var. imbricata 3 1 6 6 4sp. 4 8 8 8

Baeckia sp. # 1 1 1Bambusa

balcooa 32 50 42 5 1 3 43sp. # 1 1 1

Bambuseae sp. 8 9 7 1 1 3Banksia

ericifolia # 1 1 1integrifolia # 1 1 1 1

Bauhiniapurpurea 1 1 1sp. 1 2 2variegata 9 11 11 1

Begonia cucullata # 1 2 1 1Bidens

bipinnata * 23 23 23 7biternata * 1 1 1pilosa * 39 3 65 17 1 47 12

Billardiera heterophylla # 1 1 1Boerhavia erecta * 1 1 1Briza maxima * 1 1 1Bromus

catharticus * 2 2 1 1 1diandrus * 1 1 1 1pectinatus * 1 1 1 1

Brugmansia ×candida 6 7 1 6 1Bryophyllum delagoense 4 1 6 6?Buddleja madagascariensis # 1 1 1Caesalpinia

decapetala 127 41 413 239 25 174 153gilliesii 18 19 6 3 8 2 2

Callistemoncitrinus # 1 1 1glaucus # 1 1 1rigidus 1 1 1sp. 1 1 1 1viminalis 1 1 1 1

Calotropis procera # 1 1 1Campuloclinium macrocephalum 14 5 25 16 1 9 3Canna

glauca # 1 1 1 1indica 26 1 34 19 7 6 8 11sp. 13 17 10 4 1 3 10×generalis 7 8 8 1 5

Capsella bursa-pastoris * 1 1 1Cardiospermum

grandifl orum 43 12 57 54 7 3 29

APPENDIX 4.—Summary of results for all naturalized and casual alien plants in the study area, Savanna Biome, Fynbos Biome, Forest habitats, Grassland Biome, Nama-Karoo Biome, Succulent Karoo Biome and watercourse/wetland habitats (cont.)


records










Bothalia 37,2 (2007) 231

Cardiospermum (cont.)grandifl orum/halicacabum 16 5 20 18 4 2 5halicacabum 27 35 34 5 1 26

Carica papaya 6 6 6 2Castanea dentata ?# 1 1 1 1Castanospermum australe ?# 1 1 1Casuarina

cunninghamiana 9 12 8 2 2 8cunninghamiana/equisetifolia 42 2 63 46 6 1 11 23equisetifolia 24 1 57 55 6 2 7

Catharanthus roseus 38 1 53 49 1 2 3 11Cedrus deodara ?# 5 5 5Celtis

australis † ? ? ?occidentalis † ? ? ?sinensis † 1 1 1 ?

Cenchrus brownii * 1 1 1Centranthus ruber ?# 2 2 2 2Cereus jamacaru 124 11 193 169 6 1 12 6 6Cestrum

aurantiacum 8 1 10 7 1 6 2 2aurantiacum/laevigatum 7 3 8 5 4 3 1elegans 2 2 2 2 2laevigatum 70 16 167 138 9 30 20 41parqui 3 5 4 1sp. 1 1 1 1

Chamaesyceprostrata * 3 3 1 2serpens * 1 1 1

Chenopodium album * 3 3 3Chorizema cordatum # 1 1 1Chromolaena odorata 93 64 558 529 77 29 220Cichorium intybus 12 1 16 14 2 2Cinnamomum camphora 10 1 19 13 3 14 3 3Cirsium

arvense * 2 2 2vulgare 188 20 345 68 8 253 16 40

Citruslimon 1 1 1sp. 5 5 1 1 4 2

Coix lacryma-jobi * 1 2 1 1Colocasia esculenta 11 3 19 14 5 1 19Commelina benghalensis * 9 15 7 3 5 3 2 1Convolvulus arvensis 23 1 23 4 4 11 4 2Conyza

bonariensis * 4 4 1 3 1canadensis * 3 3 2 1 1primulifolia * 1 1 1sp.* 3 3 2 1 2sumatrensis * 1 1 1

Coreopsis lanceolata 11 15 11 4Cortaderia

jubata 7 7 2 5 1jubata/selloana 23 2 28 11 1 16 1 4selloana 54 1 104 15 77 5 10 1 1 21

Corymbia fi cifolia ?# 3 3 1 2Cosmos bipinnatus * 48 10 122 3 119 2Cotoneaster

coriaceus ?# 1 1 1franchetii 7 1 7 2 1 5 1glaucophyllus 2 2 1 1franchetii/pannosus 19 27 1 2 1 23 1 3pannosus 25 30 2 1 25 2 3



records









as ‘naturalized’ (Pyšek et al. 2004).*, mainly herbaceous species that are suspected of being under-estimated in this survey.†, Celtis australis (probably naturalized), C. occidentalis (probably naturalized) and C. sinensis (naturalized) easily mistaken for indigenous C.

africana and suspected of being under-estimated in this survey.QDSp, quarter-degree squares present; QDSa, quarter-degree squares abundant.Bold numbers in biome categories add up to upper 80% or more of total records.

232 Bothalia 37,2 (2007)

Cotoneaster (cont.)sp. 21 23 1 1 21 4

Crataegussp. # 2 2 2×lavallei 4 4 1 4

Crotalaria agatifl ora subsp. agatifl ora

18 29 24 8 1

Cryptomeria japonica # 1 1 1 1Cryptostegia grandifl ora 1 3 3 3Cuphea ignea # 1 1 1Cupressus

arizonica 47 68 1 62 5 1lusitanica 2 2 1 1 1sp. 18 24 3 1 1 19 1 2

Cuscutacampestris 82 1 103 22 3 2 73 5 23campestris/suaveolens 34 40 22 6 7 6 6 5suaveolens 7 7 2 2 4 1 1

Cydonia oblonga 7 7 3 3 1 1Cytisus scoparius 10 15 1 14 1Dahlia imperialis ?# 1 2 2Datura

ferox 175 14 201 87 3 2 85 26 55innoxia 29 3 36 30 1 1 3 2 18sp. 84 1 110 41 1 65 2 1 7stramonium 286 24 373 131 19 6 174 48 1 73

Delonix regia 5 5 5 2Desmanthus virgatus * 1 1 1Dracocephalum canariense 1 1 1Duranta erecta 32 35 33 8 2 8Dysphania ambrosioides * 2 2 1 1 1Echinopsis spachiana 57 2 83 29 2 14 37 1Echium

plantagineum 51 6 64 11 31 19 3 4plantagineum/vulgare 19 2 17 1vulgare 29 2 31 1 9 21 2

Egeria densa 2 2 2 1Eichhornia crassipes 87 72 431 279 46 106 431Eragrostis pilosa * 1 1 1Eriobotrya japonica 3 3 1 2 1 1Eucalyptus

camaldulensis 121 22 182 36 81 3 17 14 34 137cinerea 11 13 2 11cladocalyx 37 4 83 2 71 10 26cloeziana 1 1 1conferruminata 41 10 117 10 106 1 12diversicolor 49 8 153 1 148 28 2 2 32?exserta 1 1 1 1 1fastigata 1 1 1globulus 12 16 16 5 1gomphocephala 6 11 11grandis 100 16 190 111 1 19 78 67leucoxylon ?# 2 2 2 2microcorys 1 2 2microtheca 1 1 1 1paniculata 1 1 1regnans 6 7 7 3robusta ?# 1 1 1 1sideroxylon ?# 1 1 1sp. 505 30 1 103 299 147 17 614 27 16 266tereticornis ?# 1 1 1 1

Eugenia unifl ora 2 2 2Euphorbia

heterophylla 3 3 2 1 2peplus * 1 1 1pulcherrima ?# 6 9 9 1



records










Bothalia 37,2 (2007) 233

Fallopia convolvulus * 1 1 1Ficus

carica 18 26 1 19 2 1 3 13elastica # 1 1 1macrophylla # 1 1 1pumila 2 2 1 1

Flaveria bidentis * 12 12 11 1 1 6Foeniculum vulgare * 9 10 3 5 1 1 1Fraxinus

americana 13 15 11 4 3angustifolia 3 3 1 1 1sp. 13 14 11 3 4

Fuchsia sp. # 1 1 1 1Genista monspessulana 2 2 2Glandularia

aristigera * 14 27 1 26 1×hybrida ?# 1 1 1

Glebionis coronaria 2 1 4 2 2Gleditsia triacanthos 111 1 162 12 1 136 12 1 41Gnaphalium luteoalbum * 1 1 1 1Gomphrena celosioides * 2 2 2 1Grevillea

robusta 53 80 68 2 9 10 22rosmarinifolia # 1 1 1sericea # 1 1 1

Guilleminea densa * 2 2 2Hakea

drupacea 28 2 58 4 53 1gibbosa 18 3 34 2 31 1salicifolia 5 1 5 2 3 1sericea 77 17 230 14 204 3 1 11 5victoriae # 1 1 1

Harrisia martinii 21 10 33 32 1 3Hedychium

coccineum 3 1 6 6 2 2coronarium 14 1 19 18 2 1 2fl avescens 5 2 5 3 2 1 2gardnerianum 12 18 11 4 7 1sp. 7 2 8 4 3 2 1 4

Helianthus annuus * 5 8 8Heliotropium amplexicaule * 2 3 2 1Hibiscus trionum * 2 2 2Homalanthus populifolius 2 2 2 2Hordeum murinum * 1 1 1 1Hylocereus undatus 8 8 7 1Hypericum

patulum 1 1 2 1 2 1perforatum 13 1 19 16 3 3

Hypochaeris radicata * 1 1 1Ipomoea

alba 22 1 32 28 5 4 19carnea subsp. fi stulosa 21 1 36 35 1 7indica 23 3 27 22 3 3 2 8indica/purpurea 74 7 120 86 18 16 16 34nil 1 1 1 1purpurea 37 3 46 23 10 9 13 12sp. 3 3 2 1 1

Jacaranda mimosifolia 195 16 613 497 21 115 1 105Jasminum

humile 2 3 3mesnyi ?# 1 1 1

Jatrophagossypiifolia 4 10 10 1sp. 2 2 1 1



records










234 Bothalia 37,2 (2007)

Juniperuspinchotii # 1 1 1sp. 4 5 5 3virginiana 17 2 28 1 27 8

Lactuca serriola * 1 1 1Lagerstroemia indica 6 7 6 1Lantana camara 247 116 2 111 1 843 60 63 207 1 289Lemna

gibba 3 2 3 2 1 3sp. 3 1 3 1 1 1 3

Lepidiumdidymum * 1 1 1draba 4 4 1 3

Leptospermum laevigatum 38 15 102 6 96 3 10Leucaena leucocephala 36 3 123 115 11 8 43Ligustrum

japonicum 7 8 2 2 6 4lucidum 12 1 16 5 2 11 6ovalifolium 3 1 3 2 1 1 1sinense 8 11 1 10 5sp. 12 2 13 6 1 7 7vulgare 3 1 5 3 2 5

Lilium formosanum 15 3 30 18 12Limonium sinuatum 10 1 10 5 4 1 2Linaria

genistifolia * 1 1 1maroccana * 1 3 3

Litsea glutinosa 8 3 10 10 6 5Lonicera japonica var. halliana 5 5 2 1 3 4Lygodium japonicum # 1 1 1 1Lythrum salicaria 1 1 1 1Macfadyena unguis-cati 22 9 52 47 13 5 14Maireana brevifolia ?# 1 1 1Malus pumila var. paradisi-

aca ?# 5 7 2 5 1

Malvadendromorpha 16 1 24 1 19 1 3 3linnaei * 1 1 1parvifl ora * 2 2 1 1

Malvastrum coromandelianum * 2 2 1 1Mangifera indica 12 1 30 30 1 16Manihot

esculenta 8 10 9 1 1grahamii # 6 7 7 1 1

Medicago sativa * 2 2 1 1Melaleuca

hypericifolia 1 2 2wilsonii # 1 1 1

Melia azedarach 551 65 2 119 1 394 82 29 588 49 6 674Melilotus alba * 15 26 2 24 2Metasequioa glyptostroboides # 1 1 1 1Metrosideros excelsa 2 1 5 5 3Mimosa

pigra 6 1 8 7 1 7pudica var. hispida * 2 2 2

Mirabilis jalapa * 7 7 3 4Momordica charantia * 1 1 1Monstera deliciosa # 1 1 1 1Montanoa hibiscifolia 24 2 46 41 6 5 13Moringa oleifera # 2 2 2Morus alba 129 8 304 192 2 10 109 1 164Musa sp. # 8 15 15 10Myoporum tenuifolium subsp.

montanum30 49 2 44 1 3 4



records










Bothalia 37,2 (2007) 235

Myriophyllumaquaticum 48 10 81 39 17 25 81spicatum 20 23 11 2 9 1 23

Nassellatenuissima 1 2 2 1trichotoma 12 3 16 12 1 10 1 1

Nasturtium offi cinale 50 1 64 19 4 1 38 3 64Nephrolepis exaltata 13 1 19 12 6 13 1 2Nerium oleander 23 2 46 16 21 1 4 4 36Nicandra physalodes * 1 1 1Nicotiana

glauca 383 14 957 274 168 1 72 206 237 441tabacum ?# 3 3 3 3

Nymphaea×marliacea 1 1 1 1mexicana 2 2 4 1 3 4

Oenotherabiennis * 19 1 19 5 13 1 10glazioviana * 1 1 1 1indecora * 1 1 1jamesii * 15 2 17 7 1 9 10laciniata * 1 1 1rosea * 4 4 4 1sp. 4 4 4tetraptera * 1 1 1

Olyra latifolia * 1 1 1 1Opuntia

aurantiaca 61 3 84 50 28 4 2 4engelmannii (= O. lindheimeri) 10 3 15 6 2 7 1exaltata 6 6 2 3fi cus-indica 861 57 2 445 1 159 267 8 570 368 81 129fulgida 11 2 12 7 2 3humifusa 25 3 32 18 2 10 2 2?humifusa/engelmannii 48 4 49 23 17 9 1imbricata 131 15 151 49 8 3 48 43 3 6microdasys 9 10 4 3 3monacantha 48 1 114 90 22 5 2 17robusta 225 2 337 61 13 120 130 13 4sp. 37 39 21 5 10 3 2spinulifera ?# 1 1 1stricta 106 14 193 168 1 12 9 3 4?stricta × humifusa 1 1 1

Orobanche minor 4 5 2 3Oxalis corniculata * 4 4 1 3 1Pandanus sp. # 1 1 1 1Paraserianthes lophantha 54 9 286 10 274 7 2 82Parkinsonia aculeata 15 18 12 6 4Parthenium hysterophorus 15 3 29 29 8Parthenocissus quinquefolia # 1 1 1Paspalum

dilatatum * 6 6 1 2 2 3 1quadrifarium * 1 1 1 1urvillei * 1 1 1

Passifl oracaerulea 12 20 10 9 2 1 12edulis 32 55 40 6 19 9 12sp. 19 1 22 13 6 11 3 7suberosa 6 7 6 2 1subpeltata 21 1 26 22 3 4 3tripartita var. mollissima 4 4 3 1 1 1

Pennisetumclandestinum 48 12 56 12 28 3 14 2 6purpureum 40 6 87 82 5 30setaceum 66 17 84 39 20 8 17 2setaceum/villosum 15 1 15 4 4 6 1 1sp. 11 4 15 11villosum 22 5 26 2 3 14 7 1



records










236 Bothalia 37,2 (2007)

Pereskia aculeata 44 8 102 91 4 39 7 8Persea americana # 2 2 2 1Persicaria lapathifolia * 1 1 1 1Phoenix

canariensis 3 3 2 1 3dactylifera 4 4 3 1 4

Phormium tenax # 2 2 2 2Physalis

peruviana * 3 3 2 1viscosa * 3 3 1 2 2

Phytolaccadioica 30 38 17 20 1 1 15icosandra * 4 5 5 1

Pinuscanariensis 6 1 9 2 7elliottii 34 6 59 35 3 23 1 7elliottii/taeda 30 1 59 41 2 18 9halepensis 85 3 136 45 53 33 4 1 9patula 85 13 238 90 18 148 59pinaster 85 44 401 34 355 29 4 8 36pinea 18 35 1 30 4 1radiata 70 15 206 4 186 20 10 6 16roxburghii 2 2 2sp. 126 14 169 47 47 3 70 3 2 18taeda 7 11 6 5 3

Pistia stratiotes 24 6 63 60 2 1 63Pittosporum undulatum 3 7 7 1 3Pityrogramma calomelanos * 1 1 1 1Plantago

lanceolata * 4 4 4 1major * 2 2 1 1virginica * 1 1 1

Platanus sp. # 1 1 1Plectranthus comosus 19 22 8 11 1 3 2Polygonum aviculare * 1 1 1 1Polypogon monspeliensis * 1 1 1Pomaderris kumeraho # 1 1 1Pontederia cordata 2 2 2 2Populus

alba 15 2 22 11 1 10 17alba/×canescens 185 47 460 171 1 8 283 5 336deltoides 100 6 169 37 3 1 114 15 117nigra var. italica 90 120 2 7 94 17 100×canescens 371 130 939 76 279 4 486 67 31 823

Portulaca oleracea * 2 2 1 1Prosopis

glandulosa var. torreyana 40 10 50 13 2 3 29 3 13glandulosa/velutina 390 78 1 108 168 63 54 666 156 443velutina 48 6 53 7 3 1 41 1 18

Prunusarmeniaca 32 44 1 6 34 2 1 9persica 319 1 728 115 65 1 530 15 3 148serotina 1 1 1 1

Psidiumcattleianum 5 8 7 4 1 1guajava 160 50 732 662 9 25 60 1 179guineense 2 2 2 1sp. 36 7 47 44 6 3 12×durbanensis 2 3 3 2

Pterocarya stenoptera # 1 1 1 1 1Pueraria montana var. lobata 3 3 2 1 2Punica granatum 8 11 1 4 3 3 2Pyracantha

angustifolia 142 3 285 13 6 2 256 10 78angustifolia/crenulata 40 3 51 5 2 1 44 5



records










Bothalia 37,2 (2007) 237

Pyracantha (cont.)coccinea 6 7 7 2crenulata 22 31 2 1 29 2

Pyrus sp. ?# 5 5 4 1 1Quercus

canariensis ?# 1 1 1cerris ?# 2 2 2palustris 5 5 2 3 1robur 50 2 88 3 57 5 26 2 53sp. 4 6 1 4 1 3suber ?# 1 1 1

Richardiabrasiliensis * 1 1 1 1humistrata * 1 1 1

Ricinus communis 456 56 1 701 1 230 250 30 166 24 31 582Rivina humilis 7 1 10 10 4 1Robinia pseudoacacia 110 14 178 13 6 1 145 14 66Rosa

multifl ora 5 5 3 2 1? ×odorata # 1 1 1 1rubiginosa 119 12 276 12 6 2 255 3 59sp. # 3 3 2 1

Rubuscuneifolius 75 35 236 45 6 191 71fl agellaris 3 4 4 1fruticosus 89 32 244 24 188 14 24 8 71?pascuus 3 2 3 2 1phoenicolasius 4 4 1 4rosifolius 14 1 14 9 2 3 3 3sp. 86 30 179 88 5 12 86 54×proteus 4 3 4 2 2 3

Rumexacetosella subsp. pyrenaicus * 1 1 1 1crispus * 2 2 1 1usambarensis 4 1 4 3 1

Saccharum offi cinarum 15 1 26 25 1 8Salix

babylonica 475 89 1 381 140 74 6 1 069 90 8 1 323caprea 9 12 12 10fragilis 75 24 176 5 1 169 1 175

Salsola kali/tragus 155 31 187 27 13 23 117 7 2Salvinia molesta 29 7 44 25 14 5 44Sambucus

canadensis 3 1 3 1 2 3sp. 10 11 2 9 6

Scheffl era actinophylla # 1 1 1Schinus

molle 231 2 407 82 49 1 73 156 47 136terebinthifolius 30 2 90 85 2 9 3 54

Schizolobium parahyba var. parahyba #

1 1 1

Schkuhria pinnata * 4 4 1 3 1Senna

bicapsularis 16 1 45 45 31corymbosa 4 4 2 2 1didymobotrya 139 29 339 261 3 15 75 115hirsuta 9 10 10 1multiglandulosa 11 1 12 6 4 1 2 3obtusifolia 4 5 5 3occidentalis 56 4 75 74 1 1 27pendula var. glabrata 19 2 21 19 2 2 6septemtrionalis 63 102 84 1 8 17 31sp. 16 23 12 6 2 5 12

Sesbaniabispinosa * 1 1 1 1punicea 323 68 830 405 175 8 238 4 8 500



records










238 Bothalia 37,2 (2007)

Sigesbeckia orientalis * 1 1 1Silybum marianum * 7 9 7 2Sisymbrium orientale * 1 1 1Sisyrinchium sp. * 2 2 2 2Solanum

betaceum 4 5 4 3 1capsicoides * 1 1 1chrysotrichum 21 21 21 12elaeagnifolium 51 11 60 18 3 21 16 2 4mauritianum 265 99 1 364 748 86 97 530 419pseudocapsicum * 6 10 1 8 7 1 5seaforthianum 30 3 77 75 1 31 1 50sisymbriifolium 40 3 51 18 3 1 30 5sp. 4 6 5 2 1 1torvum * 1 1 1

Sonchus oleraceus * 3 3 1 2Sophora cf. davidii # 1 1 1 1Sorghum halepense 41 2 46 26 3 2 11 6 17Spartium junceum 20 3 43 4 35 3 1 2Spathodea campanulata 3 4 4Sphagneticola trilobata * 1 2 2 1Spiraea cantoniensis # 1 1 1Stellaria media * 1 1 1Stenocarpus sinuatus ?# 1 1 1 1Styphnolobium japonicum ?# 1 1 1 1Symphyotrichum squamatum * 1 1 1Syncarpia glomulifera 2 2 2Syzygium

cumini 9 1 14 13 2 1 4jambos 3 3 2 1 2paniculatum 3 3 3

Tabebuia chrysotricha ?# 1 1 1Tagetes minuta * 47 78 11 1 66 1 16Tamarix

chinensis 4 4 1 3 2ramosissima 7 8 1 3 4 8sp. 85 4 110 10 16 8 58 18 85

Taraxacum offi cinale * 1 1 1Tecoma stans 57 4 99 95 2 4 17Tephrocactus

?aoracanthus 1 1 1 1articulatus 1 1 1sp. 1 1 1

Thevetia peruviana 15 23 23 6Tipuana tipu 24 1 42 33 9 3Tithonia

diversifolia 49 5 123 120 10 3 33diversifolia/rotundifolia 3 1 3 3rotundifolia 19 5 28 22 2 6 7

Toona ciliata 23 1 54 51 7 3 10Torilis arvensis * 1 1 1Toxicodendron succedaneum 12 1 14 12 3 2 3Tragopogon dubius * 1 1 1Tridax procumbens * 1 1 1 1Triplaris americana ?# 1 1 1Triticum aestivum * 1 1 1Tropaeolum majus * 3 3 2 1 1 1Ulex europaeus 9 14 2 11 1 3Ulmus

parvifolia 2 2 2 1procera # 1 1 1 1sp. # 4 4 3 1 1

Verbenabonariensis * 58 4 115 22 93 13brasiliensis/bonariensis 2 4 3 1brasiliensis * 2 2 2



records










Bothalia 37,2 (2007) 239



Verbena (cont.)offi cinalis * 2 2 2 1rigida var. rigida * 1 1 1

Verbesina encelioides * 18 21 14 7 5Vinca major * 1 1 1Vitis sp. # 6 12 11 1Washingtonia sp. ?# 3 3 2 1 3Wigandia urens var.

caracasana # 3 4 2 2

Wisteria fl oribunda # 1 1 1Xanthium

sp. 6 7 5 2 1 7spinosum 83 6 104 30 3 62 8 27strumarium 149 21 212 126 3 4 72 11 95

Yucca aloifolia 20 22 8 2 9 2 5Zinnia peruviana * 4 6 5 1 1

Acaciabaileyana F.Muell., Fabaceae, Bailey’s wattlecultriformis A.Cunn. ex G.Don, Fabaceae, knife-leaved wattle #cyclops A.Cunn. ex G.Don, Fabaceae, red eyedealbata Link., Fabaceae, silver wattledecurrens Willd., Fabaceae, green wattleelata A.Cunn. ex Benth. (A. terminalis (Salisb.) J.F.Macb. misapplied

in South Africa), Fabaceae, peppertree wattlefi mbriata A.Cunn. ex G.Don, Fabaceae, fringed wattleimplexa Benth., Fabaceae, hickory wattlelongifolia (Andrews) Willd., Fabaceae, long-leaved wattlemearnsii De Wild., Fabaceae, black wattlemelanoxylon R.Br., Fabaceae, Australian blackwoodparadoxa DC. (= A. armata R.Br.), Fabaceae, kangaroo thornpodalyriifolia A.Cunn. ex G.Don, Fabaceae, pearl acaciapycnantha Benth., Fabaceae, golden wattlesaligna (Labill.) H.L.Wendl. (= A. cyanophylla Lindl.), Fabaceae, Port

Jackson willowstricta (Andrews) Willd., Fabaceae, hop wattle ?#, *2004 viscidula Benth., Fabaceae, sticky wattle

Acanthocereus ?tetragonus (L.) Hummelinck, Cactaceae, barbed-wire cactus

Acanthospermumaustrale (Loefl .) Kuntze (= A. brasilum Schrank), Asteraceae, eight-

seeded prostrate starburhispidum DC., Asteraceae, upright starbur

Acanthus polystachyus Delile var. pseudopubescens Cufod. (= A. pubescens Engl.), Acanthaceae, bear’s breeches #

Acerbuergerianum Miq., Aceraceae, Chinese maple, *2003 #negundo L. (= A. californicum D.Dietr.), Aceraceae, ash-leaved maple? sp., Aceraceae, ?red-leafed maple

Achyranthes aspera L. (= A. argentea Lam.), Amaranthaceae, burweedAcorus calamus L., Acoraceae, calamusAcrocarpus fraxinifolius Wight ex Arn., Fabaceae, shingle tree #Adiantum raddianum C.Presl, Adiantaceae, maidenhair fern #Agave

americana Lvar. americana, Agavaceae, American agave

var. expansa (Jacobi) Gentry (= A. expansa Jacobi), Agavaceae, spreading century plant

decipiens Baker (= A. laxifolia Baker), Agavaceae, false sisal #sisalana Perrine, Agavaceae, sisalsp., Agavaceae

Ageratinaadenophora (Spreng.) R.M.King & H.Rob. (= Eupatorium adenophorum

Spreng.), Asteraceae, crofton weedriparia (Regel) R.M.King & H.Rob. (= Eupatorium riparium Regel),

Asteraceae, creeping crofton weed ?#Ageratum

conyzoides L., Asteraceae, invading ageratumhoustonianum Mill., Asteraceae, Mexican ageratum

Agrimonia cf. parvifl ora Aiton, Rosaceae, agrimony #Agrostemma githago L., Caryophyllaceae, corn cockleAilanthus altissima (Mill.) Swingle, Simaroubaceae, tree-of-heavenAlbizia

chinensis (Osbeck) Merr. (= A. stipulata (DC.) Boivin), Fabaceae, Chinese false-thorn #

lebbeck (L.) Benth., Fabaceae, lebbeck treeprocera (Roxb.) Benth., Fabaceae, false lebbeck

Alhagi maurorum Medik. (= A. camelorum Fisch.), Fabaceae, camelthorn bush

Alisma plantago-aquatica L., Alismataceae, water plantainAlnus glutinosa (L.) Gaertn. (= A. barbata C.A.Mey.), Betulaceae,

black elderAlpinia zerumbet (Pers.) B.L.Burtt & R.M.Sm. (= A. speciosa (J.C.Wendl.)

K.Schum.), Zingiberaceae, shell gingerAlternanthera pungens Kunth (= A. repens (L.) Link.), Amaranthaceae,

khaki bur weedAmaranthus

hybridus L. , Amaranthaceae, pigweedsp., Amaranthaceae

Ambrosia artemisiifolia L., Asteraceae, annual ragweedAmmi majus L. (= A. glaucifolium L.), Apiaceae, bishop’s weedAnigozanthos fl avidus DC., Haemodoraceae, yellow kangaroo paw #Anredera cordifolia (Ten.) Steenis, (A. baselloides (Kunth) Baill.

misapplied in South Africa), Basellaceae, bridal wreath

APPENDIX 5.—Species checklist

The following 601naturalized and casual alien (#) plant species were catalogued in the SAPIA database up to May 2006. Accepted names in roman type. Synonyms in italics. *, taxa added to SAPIA after 2000; †, taxa recorded only in Zimbabwe and Malawi. PRE, species records from the Pretoria National Herbarium



records








240 Bothalia 37,2 (2007)

Antigonon leptopus Hook. & Arn., Polygonaceae, coral creeperApium graveolens L., Apiaceae, wild celeryAraucaria

sp., Araucariaceae, monkey puzzle tree, *2003 #bidwillii Hook., Araucariaceae, bunya-bunya, *2005 #

Araujia sericifera Brot., Asclepiadaceae, moth catcherArdisia

crenata Sims (A. crispa (Thunb.) A.DC. misapplied in South Africa), Myrsinaceae, coralberry tree

elliptica Thunb. (= A. humilis Vahl), Myrsinaceae, shoebutton ardisia, *2005 #

Argemonemexicana L., Papaveraceae, yellow-fl owered Mexican poppyochroleuca Sweet subsp. ochroleuca, Papaveraceae, white-fl owered

Mexican poppysp., Papaveraceae

Aristolochia elegans Mast., Aristolochiaceae, calico fl owerArundo donax L., Poaceae, giant reedAstartea fascicularis (Labill.) DC., Myrtaceae #Atriplex

infl ata F.Muell. (= A. lindleyi Moq. subsp. infl ata (F.Muell.) Paul G.Wilson), Chenopodiaceae, sponge-fruit saltbush

muelleri Benth., Chenopodiaceae, Mueller’s saltbushnummularia Lindl. subsp. nummularia, Chenopodiaceae, old-man

saltbushsemibaccata R.Br., Chenopodiaceae, Australian saltbushsp., Chenopodiaceae

Azollafi liculoides Lam., Azollaceae, red water fern?pinnata R.Br. subsp. asiatica R.M.K Saunders & K.Fowler (= A.

imbricata (Roxb. ex Griff.) Nakai), Azollaceae, mosquito fernsp., Azollaceae

Baeckia sp., Myrtaceae #Bambusa

balcooa Roxb., Poaceae, common bamboosp. with tall yellow stems and green leaves, Poaceae, bamboo

Bambuseae sp., Poaceae, bambooBanksia

ericifolia L.f., Proteaceae, heath banksia #integrifolia L.f., Proteaceae, coast banksia #

Bauhiniapurpurea L., Fabaceae, butterfl y orchid treesp., Fabaceaevariegata L., Fabaceae, orchid tree

Begonia cucullata Willd. (= B. semperfl orens Link & Otto), Begoniaceae, begonia #

Bidensbipinnata L., Asteraceae, Spanish black jackbiternata (Lour.) Merr. & Sherff, Asteraceae, fi ve-leaved black jackpilosa L., Asteraceae, black jack

Billardiera heterophylla (Lindl.) L.W.Cayzer & Crisp (= Sollya heterophylla Lindl.), Pittosporaceae, bluebell creeper #

Boerhavia erecta L., Nyctaginaceae, erect boerhaviaBougainvillea glabra Choisy, Nyctaginaceae, bougainvillea, *2004 #Brachychiton populneus (Schott & Endl.) R.Br., Sterculiaceae,

kurrajong, *2006 # Briza maxima L. (= B. major K.Presl), Poaceae, quaking grassBromus

catharticus Vahl (= B. unioloides Kunth, B. willdenowii Kunth), Poaceae, rescue grass

diandrus Roth, Poaceae, ripgut bromepectinatus Thunb. (= B. adoensis Hochst. ex Steud.), Poaceae,

Japanese bromeBrugmansia ×candida Pers., (= Datura candida (Pers.) Saff.), Solan-

aceae, moonfl ower bushBryophyllum

delagoense (Eckl. & Zeyh.) Schinz (= Kalanchoe tubifl ora (Harv.) Raym.-Hamet), Crassulaceae, chandelier plant

pinnatum (Lam.) Oken (= Kalanchoe pinnata (Lam.) Pers.), Crassu-laceae, green mother of millions, *2005 ?naturalized

proliferum Bowie ex Hook. (= Kalanchoe prolifera (Bowie ex Hook.) Raym.-Hamet, Crassulaceae, *2005 ?naturalized

Buddlejadavidii Franch., Buddlejaceae, Chinese sagewood, *2004 ?naturalized

?Buddleja madagascariensis Lam., Buddlejaceae, Madagascar sage-wood #

Caesalpiniadecapetala (Roth) Alston (= C. sepiaria Roxb.), Fabaceae, Mauritius

thorngilliesii (Hook.) D.Dietr., Fabaceae, bird-of-paradisepulcherrima (L.) Sw., Fabaceae, pride of barbados, *2004 #

Callisia repens (Jacq.) L., Commelinaceae, creeping inch plant, *2006 # Callistemon

citrinus (Curtis) Skeels, Myrtaceae, crimson bottlebrush #glaucus (Curtis) Sweet (= C. speciosus auct.), Myrtaceae, Albany

bottlebrush #rigidus R.Br., Myrtaceae, stiff bottlebrushsp., Myrtaceaeviminalis (Sol. ex Gaertn.) G.Don, Myrtaceae, weeping bottlebrush

Calotropis procera (Aiton) W.T.Aiton (= Asclepias procera Aiton), Ascle-piadaceae, madar #

Campuloclinium macrocephalum (Less.) DC. (= Eupatorium macro-cephalum Less.), Asteraceae, pom pom weed

Cannaglauca L., Cannaceae, yellow-fl owered glaucous canna #indica L. (= C. edulis Ker Gawl.), Cannaceae, Indian cannasp., Cannaceae×generalis L.H.Bailey, Cannaceae, garden canna

Capsella bursa-pastoris (L.) Medik., Brassicaceae, shepherd’s purseCardiospermum

grandifl orum Sw., Sapindaceae, balloon vinehalicacabum L., Sapindaceae, heart pea

Carica papaya L. (= Papaya carica Gaertn.), Caricaceae, pawpawCastanea dentata (Marshall) Borkh., Fagaceae, American chestnut ?#Castanospermum australe A.Cunn. & C.Fraser ex Hook., Fabaceae,

Australian chestnut ?naturalizedCasuarina

cunninghamiana Miq., Casuarinaceae, beefwoodequisetifolia L., Casuarinaceae, horsetail tree

Catharanthus roseus (L.) G.Don (= Lochnera rosea (L.) Rchb., Vinca rosea L.), Apocynaceae, Madagascar periwinkle

Cedrus deodara (Roxb. ex D.Don) G.Don, Pinaceae, deodar ?#Celtis

australis L., Ulmaceae, European hackberry ?naturalizedoccidentalis L., Ulmaceae, common hackberry ?naturalizedsinensis Pers., Ulmaceae, Chinese nettle tree

Cenchrus brownii Roem. & Schult. (= C. viridis Spreng.), Poaceae, fi ne burgrass

Centranthus ruber (L.) DC., Valerianaceae, red valerian ?naturalizedCereus jamacaru DC. (C. peruvianus (L.) Mill. misapplied in South

Africa), Cactaceae, queen of the nightCestrum

aurantiacum Lindl., Solanaceae, yellow or orange cestrumelegans (Brongn.) Schltdl. (= C. purpureum (Lindl.) Standl.), Solan-

aceae, crimson cestrumlaevigatum Schltdl., Solanaceae, inkberryparqui L’Hér., Solanaceae, Chilean cestrumsp., Solanaceae

Chamaesyceprostrata (Aiton) Small (= Euphorbia prostrata Aiton), Euphorbiaceae,

hairy creeping milkweedserpens (Kunth) Small (= Euphorbia serpens Kunth), Euphorbiaceae,

milkweedChenopodium album L., Chenopodiaceae, white goosefootChondrilla juncea L., Asteraceae, skeletonweed, *2003 #Chorizema cordatum Lindl., Fabaceae, Australian fl ame pea #Chromolaena odorata (L.) R.M.King & H.Rob. (= Eupatorium odoratum

L.), Asteraceae, triffi d weedCichorium intybus L., Asteraceae, chicoryCinnamomum camphora (L.) J.Presl, Lauraceae, camphor treeCirsium

arvense (L.) Scop., Asteraceae, Canada thistlevulgare (Savi) Ten. (= C. lanceolatum (L.) Scop.), Asteraceae, spear

thistleCissus antarctica Vent., Vitaceae, kangaroo vine, *2002 #Citrus

limon (L.) Burm.f. (= C. limonum Risso), Rutaceae, lemonsp., Rutaceae

Clusia rosea Jacq., Clusiaceae, balsam fi g/apple, *2003 #Coffea arabica L., Rubiaceae, arabica coffee, †Zimbabwe # abundant

locallyCoix lacryma-jobi L., Poaceae, Job’s tearsColocasia esculenta (L.) Schott, Araceae, elephant’s ear

APPENDIX 5.—Species checklist (cont.)

Bothalia 37,2 (2007) 241

Commelina benghalensis L., Commelinaceae, Benghal wandering JewConvolvulus arvensis L., Convolvulaceae, fi eld bindweedConyza

bonariensis (L.) Cronquist (= Erigeron bonariensis L.), Asteraceae, fl ax-leaf fl eabane

canadensis (L.) Cronquist (= Erigeron canadensis L.), Asteraceae, horseweed fl eabane

primulifolia (Lam.) Cuatrec. & Lourteig (= C. chilensis Spreng.), Asteraceae, Chilean fl eabane

sp., Asteraceaesumatrensis (Retz.) E.Walker (= C. albida Willd. ex Spreng.),

Asteraceae, tall fl eabaneCoreopsis lanceolata L., Asteraceae, tickseedCortaderia

jubata (Lemoine ex Carrière) Stapf, Poaceae, purple Pampas grassselloana (Schult.) Asch. & Graebn., Poaceae, common Pampas grass

Corymbia fi cifolia (F.Muell.) K.D.Hill & L.A.S.Johnson (= Eucalyptus fi cifolia F.Muell.), Myrtaceae, red fl owering gum #

Cosmos bipinnatus Cav. (= Bidens formosa (Bonato) Sch. Bip.), Asteraceae, cosmos

Cotoneastercoriaceus Franch. (= C. lacteus W.W.Sm.), Rosaceae #franchetii Bois, Rosaceae, orange cotoneasterglaucophyllus Franch., Rosaceae, late cotoneasterpannosus Franch., Rosaceae, silver-leaf cotoneastersp., Rosaceae

Crataegussp., Rosaceae #×lavallei Hérincq (= C. carrierei Vauvel ex Carriére), Rosaceae,

Lavallee thornCrotalaria agatifl ora Schweinf. subsp. agatifl ora, Fabaceae, canary-bird

bushCryptomeria japonica (L.f.) D.Don, Cupressaceae, Japanese cedar #Cryptostegia grandifl ora R.Br., Asclepiadaceae, rubber vineCuphea ignea A.DC., Lythraceae, cigarette bush #Cupressus

arizonica Greene (= C. glabra Sudw.), Cupressaceae, Arizona cypresslusitanica Mill. (= C. lindleyi Klotzsch ex Endl.), Cupressaceae, Mexi-

can cypresssp., Cupressaceae

Cuscutacampestris Yunck., Convolvulaceae, common doddersuaveolens Ser., Convolvulaceae, lucerne dodder

Cyathea cooperi (Hook. ex F.Muell.) Domin (= Sphaeropteris cooperi (Hook. ex F. Muell.) R.M.Tryon), Cyatheaceae, Australian tree fern, *2005 #

Cydonia oblonga Mill.( = C. vulgaris Pers.), Rosaceae, quinceCytisus scoparius (L.) Link (= Genista scoparia (L.) Lam.), Fabaceae,

Scotch broomDahlia spp., Asteraceae, garden dahlias ?naturalizedDatura

ferox L., Solanaceae, large thorn appleinnoxia Mill. (D. metel L. misapplied in South Africa), Solanaceae,

downy thorn applesp., Solanaceaestramonium L., Solanaceae, common thorn apple

Delonix regia (Bojer ex Hook.) Raf. (= Poinciana regia Bojer ex Hook.), Fabaceae, fl amboyant

Desmanthus virgatus (L.) Willd. (= D. depressus Humb. & Bonpl. ex Willd.), Fabaceae, ground tamarind

Desmodium uncinatum (Jacq.) DC., Fabaceae, silverleaf desmodium, †Zimbabwe # abundant locally

Dracocephalum canariense L. (= Cedronella canariensis (L.) Webb & Berthel.), Lamiaceae, hortela de burro

Duranta erecta L. (= D. repens L., D. plumieri Jacq.), Verbenaceae, forget-me-not-tree

Dysphania ambrosioides (L.) Mosyakin & Clemants (= Chenopodium ambrosioides L.), Chenopodiaceae, American goosefoot

Echinopsis spachiana (Lem.) Friedrich & G.D.Rowley (= Trichocereus spachianus (Lem.) Riccob.), Cactaceae, torch cactus

Echiumplantagineum L. (= E. lycopsis L.), Boraginaceae, Patterson’s cursevulgare L., Boraginaceae, blue echium

Egeria densa Planch. (= Elodea densa (Planch.) Casp.), Hydrocharitaceae, dense water weed

Eichhornia crassipes (Mart.) Solms, Pontederiaceae, water hyacinthEragrostis pilosa (L.) P.Beauv., Poaceae, Indian love grassEriobotrya japonica (Thunb.) Lindl., Rosaceae, loquat

Eucalyptuscamaldulensis Dehnh., Myrtaceae, red river gumcinerea F.Muell. ex Benth., Myrtaceae, fl orist’s gumcladocalyx F.Muell., Myrtaceae, sugar gumcloeziana F.Muell., Myrtaceae, iron gumconferruminata D.J.Carr & S.G.M.Carr (E. lehmannii (Schauer)

Benth. misapplied in South Africa), Myrtaceae, bald island marlock or ‘spider gum’

diversicolor F.Muell., Myrtaceae, karri?exserta F.Muell., Myrtaceae, Queensland peppermintfastigata H.Deane & Maiden, Myrtaceae, cut-tail gumglobulus Labill., Myrtaceae, blue gumgomphocephala DC., Myrtaceae, tuartgrandis W.Hill ex Maiden (E. saligna Sm. misapplied in South Africa),

Myrtaceae, saligna gumleucoxylon F.Muell., Myrtaceae, white ironbark ?#microcorys F.Muell., Myrtaceae, tallow gummicrotheca F.Muell., Myrtaceae, coolabahpaniculata Sm., Myrtaceae, grey ironbarkregnans F.Muell., Myrtaceae, mountain ashrobusta Sm., Myrtaceae, swamp mahogany gum ?#sideroxylon A.Cunn ex Woolls, Myrtaceae, black ironbark ?#sp., Myrtaceaetereticornis Sm., Myrtaceae, forest red gum ?#

Eugenia unifl ora L., Myrtaceae, pitangaEuphorbia

heterophylla L. (= E. geniculata Ortega), Euphorbiaceae, annual poinsettia

leucocephala Lotsy, Euphorbiaceae, white poinsettia, *2005 ?naturalized

peplus L., Euphorbiaceae, stinging milkweedpulcherrima Willd. ex Klotzsch (= Poinsettia pulcherrima (Willd. ex

Klotzsch) Graham), Euphorbiaceae, poinsettia ?#Euryops chrysanthemoides (DC.) B.Nord., Asteraceae, †Zimbabwe #

but indigenous in South AfricaFallopia

convolvulus (L.) Á.Löve (= Bilderdykia convolvulus (L.) Dumort), Polygonaceae, climbing knotweed

sachalinensis (F.Schmidt) Ronse Decr. (= Polygonum sachalinense F.Schmidt, Reynoutria sachalinensis (F.Schmidt) Nakai), Poly-gonaceae, giant knotweed, *2005 (PRE 1980) naturalized

Ficuscarica L., Moraceae, fi gelastica Roxb. ex Hornem.(= F. decora hort.), Moraceae, rubber fi g #macrophylla Desf. ex Pers., Moraceae, Australian banyan #pumila L., Moraceae, tickey creeper

Flaveria bidentis (L.) Kuntze (= F. contrayerba (Cav.) Pers.), Asteraceae, smelter’s bush

Foeniculum vulgare Mill., Apiaceae, fennelFraxinus

americana L., Oleaceae, American ashangustifolia Vahl, Oleaceae, Algerian ashsp., Oleaceae

Fuchsia sp., Onagraceae, fuchsia #Fumaria muralis Sond. ex Koch, Fumariaceae, wall fumitory, *2001 #Genista monspessulana (L.) L.A.S.Johnson (= Cytisus candicans (L.)

DC., C. monspessulanus L.), Fabaceae, Montpellier broomGlandularia

aristigera (S.Moore) Tronc.(= Verbena tenuisecta Briq.), Verbenaceae, fi ne-leaved verbena

×hybrida (hort. ex Groenl. & Rümpler) G.L.Nesom & Pruski (= Verbena ×hybrida hort. ex Groenl. ex Rümpler), Verbenaceae, garden verbena ?#

Glebionis coronaria (L.) Cass. ex Spach (= Chrysanthemum coronarium L.), Asteraceae, chrysanthemum greens

Gleditsia triacanthos L., Fabaceae, honey locustGlyceria maxima (Hartm.) Holmb. (= G. aquatica (L.) Wahlb., Poa

aquatica L.), Poaceae, reed meadow grass, *2002 #Gmelina arborea Roxb., Verbenaceae, white teak, †Malawi # abundant

locallyGnaphalium luteoalbum L. (= Pseudognaphalium luteoalbum (L.)

Hilliard & B.L.Burtt), Asteraceae, Jersey cudweedGomphrena celosioides Mart. (= G. decumbens Jacq.), Amaranthaceae,

prostrate globe amaranthGrevillea

banksii R.Br., Proteaceae, Bank’s grevillea, *2004 # very abundant locally

robusta A.Cunn. ex R.Br., Proteaceae, Australian silky oakrosmarinifolia A.Cunn., Proteaceae #sericea (Sm.) R.Br., Proteaceae, pink spider fl ower #

Guilleminea densa (Humb. & Bonpl. ex Schult.) Moq. (= Brayulinea densa (Willd.) Small), Amaranthaceae, carrot weed


242 Bothalia 37,2 (2007)

Hakeadrupacea (C.F.Gaertn.) Roem. & Schult. (= H. suaveolens R.Br.),

Proteaceae, sweet hakeagibbosa (Sm.) Cav., Proteaceae, rock hakeasalicifolia (Vent.) B.L.Burtt (= H. saligna (Andrews) Knight), Proteaceae,

willow hakeasericea Schrad. & J.C.Wendl., Proteaceae, silky hakeavictoriae J.Drumm., Proteaceae #

Harrisia martinii (Labour.) Britton & Rose (= Eriocereus martinii (Labour.) Riccob.), Cactaceae, harrisia

Hedera helix L. subsp. canariensis (Willd.) Cout., Araliaceae, Algerian or Canary ivy, *2003 #

Hedychiumcoccineum Buch.-Ham. ex Sm., Zingiberaceae, red ginger lilycoronarium J.König, Zingiberaceae, white ginger lilyfl avescens Carey ex Roscoe, Zingiberaceae, yellow ginger lilygardnerianum Sheppard ex Ker Gawl., Zingiberaceae, kahili ginger

lilysp., Zingiberaceae

Helianthus annuus L., Asteraceae, common sunfl owerHeliotropium amplexicaule Vahl, Boraginaceae, blue heliotropeHibiscus trionum L., Malvaceae, bladderweedHomalanthus populifolius Graham, Euphorbiaceae, Queensland poplarHordeum murinum L., Poaceae, wild barleyHydrilla verticillata (L.f.) Royle, Hydrocharitaceae, hydrilla, *2006

(PRE 1963 but misidentifi ed) naturalized and very abundant at Pongolapoort Dam, KwaZulu-Natal

Hydrocotyle ranunculoides L.f., Apiaceae, †Zimbabwe # abundant locally

Hylocereus undatus (Haw.) Britton & Rose, Cactaceae, night-blooming cereus

Hypericumpatulum Thunb. (= H. patulum var. forrestii Chitt.), Clusiaceae perforatum L., Clusiaceae, St. John’s wort

Hypochaeris radicata L., Asteraceae, hairy wild lettuceHypoestes phyllostachya Baker, Acanthaceae, polka-dot-plant, *2002 #Ipomoea

alba L., Convolvulaceae, moonfl owercarnea Jacq. subsp. fi stulosa (Mart. ex Choisy) D.F.Austin (= I.

fi stulosa Mart. ex Choisy), Convolvulaceae, potato bushindica (Burm.) Merr. (= I. congesta R.Br.), Convolvulaceae, perennial

morning glorynil (L.) Roth, Convolvulaceae, Japanese morning glorypurpurea (L.) Roth, Convolvulaceae, common morning glorysp., Convolvulaceae

Iris pseudacorus L., Iridaceae, yellow fl ag, *2004 #Jacaranda mimosifolia D.Don, Bignoniaceae, jacarandaJasminum

humile L., Oleaceae, yellow bush jasminemesnyi Hance, Oleaceae, primrose jasmine ?#polyanthum Franch., Oleaceae, creeping jasmine, *2001 #

Jatrophacurcas L., Euphorbiaceae, physic nut, *2005 (1979 in Wells et al.

(1986))gossypiifolia L., Euphorbiaceae, coral plantsp., Euphorbiaceae

Juniperuspinchotii Sudw., Cupressaceae, red-berry juniper #sp., Cupressaceaevirginiana L., Cupressaceae, red cedar

Koelreuteria paniculata Laxm. (= K. apiculata Rehder & E.H.Wilson), Sapindaceae, golden-rain tree *2001 #

Lactuca serriola L. (= L. scariola L.), Asteraceae, wild lettuceLagerstroemia indica L., Lythraceae, pride-of-IndiaLantana camara L., Verbenaceae, lantanaLemna

gibba L., Lemnaceae, duckweedsp., Lemnaceae

Lepidiumdidymum L. (= Coronopus didymus (L.) Sm.), Brassicaceae,

swinecressdraba L. (= Cardaria draba (L.) Desv.), Brassicaceae, hoary cardaria

Leptospermum laevigatum (Gaertn.) F.Muell., Myrtaceae, Australian myrtle

Leucaena leucocephala (Lam.) de Wit (= L. glauca Benth.), Fabaceae, leucaena

Ligustrumjaponicum Thunb., Oleaceae, Japanese wax-leaved privetlucidum W.T.Aiton, Oleaceae, Chinese wax-leaved privetovalifolium Hassk., Oleaceae, Californian privet

sinense Lour., Oleaceae, Chinese privetsp., Oleaceaevulgare L., Oleaceae, common privet

Lilium formosanum Wallace (= L. longifl orum Thunb. var. formosanum Baker, L. phillipinense Baker), Liliaceae, Saint Joseph’s lily

Limonium sinuatum (L.) Mill. (= Statice sinuata L.), Plumbaginaceae, statice

Linariagenistifolia (L.) Mill. (= L. dalmatica (L.) Mill.), Scrophulariaceae,

yellow linariamaroccana Hook.f., Scrophulariaceae, baby snapdragon

Litsea glutinosa (Lour.) C.B.Rob. (= L. sebifera Pers.), Lauraceae, Indian laurel

Lonicera japonica Thunb. ‘Halliana’, Caprifoliaceae, Japanese honey-suckle

Lygodium japonicum (Thunb.) Sw., Schizaeaceae, Japanese climbing fern #

Lythrum salicaria L., Lythraceae, purple loosestrifeMacfadyena unguis-cati (L.) A.H.Gentry, Bignoniaceae, cat’s claw

creeperMaireana brevifolia (R.Br.) Paul G.Wilson (= Kochia brevifolia R.Br.),

Chenopodiaceae, small-leaf bluebush #Malus pumila Mill. var. paradisiaca C.K.Schneid., Rosaceae, paradise

apple ?#Malva

dendromorpha M.F.Ray (= Lavatera arborea L.), Malvaceae, tree mallow

linnaei M.F.Ray (= Lavatera cretica L.), Malvaceae, Cretan holly-hock

parvifl ora L., Malvaceae, small mallowMalvastrum coromandelianum (L.) Garcke, Malvaceae, prickly

malvastrumMangifera indica L., Anacardiaceae, mangoManihot

esculenta Crantz (= M. utilissima Pohl), Euphorbiaceae, bitter cassava

grahamii Hook. (= M. dulcis (J.F.Gmel.) Pax var. multifi da (Graham) Pax), Euphorbiaceae, hardy cassava #

Medicago sativa L. (= M. falcata L.), Fabaceae, alfalfaMelaleuca

hypericifolia Sm., Myrtaceae, red-fl owering tea treewilsonii F.Muell., Myrtaceae, violet honey-myrtle #

Melia azedarach L., Meliaceae, seringa or ‘syringa’Melilotus alba Medik., Fabaceae, white sweet cloverMetasequoia glyptostroboides Hu & W.C.Cheng, Cupressaceae, dawn

redwood #Metrosideros excelsa Sol. ex Gaertn. (= M. tomentosa A. Rich.),

Myrtaceae, New Zealand bottlebrushMichelia champaca L., Magnoliaceae, champac magnolia, †Zimbabwe #Mimosa

pigra L., Fabaceae, giant sensitive plantpudica L. var. hispida Brenan, Fabaceae, sensitive plant

Mirabilis jalapa L., Nyctaginaceae, four-o’clockMomordica charantia L., Cucurbitaceae, bitter cucumberMondia whitei (Hook.f.) Skeels (= Chlorocodon whitei Hook.f.), Apocyn-

aceae, †Zimbabwe & Malawi # but indigenous in South AfricaMonstera deliciosa Liebm., Araceae, Swiss-cheese plant #Montanoa hibiscifolia Benth., Asteraceae, tree daisyMoringa oleifera Lam. (= M. pterygosperma Gaertn.), Moringaceae,

horse-radish tree #Morus alba L., Moraceae, white or common mulberryMusa sp., Musaceae, banana tree #Murraya paniculata (L.) Jack. (= M. exotica L. ), Rutaceae, orange

jessamine, *2005 #Myoporum tenuifolium G.Forst. subsp. montanum (R.Br.) Chinnock (=

M. montanum R.Br.)(M. acuminatum R.Br. misapplied in South Africa), Myoporaceae, manatoka

Myriophyllum aquaticum (Vell.) Verdc. (= M. brasiliense Cambess.), Haloragaceae,

parrot’s featherspicatum L., Haloragaceae, spiked water-milfoil

Nassellatenuissima (Trin.) Barkworth (= Stipa tenuissima Trin.), Poaceae, white

tussocktrichotoma (Nees) Hack. ex Arechav. (= Stipa trichotoma Nees),

Poaceae, nassella tussockNasturtium offi cinale R.Br. (= Rorippa nasturtium-aquaticum (L.)

Hayek), Brassicaceae, watercressNephrolepis exaltata (L.) Schott, Nephrolepidaceae, sword fern


Bothalia 37,2 (2007) 243

Nerium oleander L., Apocynaceae, oleanderNicandra physalodes (L.) Gaertn., Solanaceae, apple-of-PeruNicotiana

glauca Graham, Solanaceae, wild tobaccotabacum L., Solanaceae, tobacco ?#

Nymphaeamexicana Zucc., Nymphaeaceae, yellow waterlily×marliacea W.Watson, Nymphaeaceae, Marliac hybrid waterlily

Oenotherabiennis L., Onagraceae, evening primroseglazioviana Micheli (= O. erythrosepala Borbás), Onagraceae, evening

primroseindecora Cambess., Onagraceae, evening primrosejamesii Torr. & A.Gray, Onagraceae, giant evening primroselaciniata Hill, Onagraceae, cutleaf evening primroserosea L’Hér. ex Aiton, Onagraceae, rose evening primrosesp., Onagraceaetetraptera Cav., Onagraceae, white evening primrose

Olyra latifolia L., Poaceae Opuntia

aurantiaca Lindl., Cactaceae, jointed cactusengelmannii Salm-Dyck ex Engelm. (= O. lindheimeri Engelm.),

Cactaceae, small round-leaved prickly pearexaltata A.Berger (= Austrocylindropuntia exaltata (A.Berger)

Backeb.), Cactaceae, long-spine cactusfi cus-indica (L.) Mill. (= O. megacantha Salm-Dyck), Cactaceae,

sweet prickly pearfulgida Engelm. (= Cylindropuntia fulgida (Engelm.) F.M.Knuth)(O.

rosea DC. and Cylindropuntia rosea (DC.) Backeb. misapplied in South Africa), Cactaceae, chainfruit cholla or ‘rosea cactus’

humifusa (Raf.) Raf. (= O. compressa auct.), Cactaceae, large-fl owered prickly pear

imbricata (Haw.) DC. (= Cylindropuntia imbricata (Haw.) F.M.Knuth), Cactaceae, imbricate prickly pear

microdasys (Lehm.) Pfeiff., Cactaceae, yellow bunny-earsmonacantha Haw. (= O. vulgaris auct.), Cactaceae, cochineal prickly

pearrobusta H.L.Wendl. ex Pfeiff., Cactaceae, blue-leaf cactussp., Cactaceaespinulifera Salm-Dyck, Cactaceae, large round-leaved prickly pear ?#stricta (Haw.) Haw. (possibly both var. dillenii and var. stricta), Cac-

taceae, Australian pest pearstricta ×humifusa?, Cactaceae tomentosa Salm-Dyck, Cactaceae, velvet opuntia, *2003 #

Orobanche minor Sm., Orobanchaceae, clover broomrapeOxalis corniculata L., Oxalidaceae, creeping oxalisPandanus sp., Pandanaceae, screw-pine #Paraserianthes lophantha (Willd.) I.C.Nielsen (= Albizia lophantha

(Willd.) Benth.), Fabaceae, stinkbeanParkinsonia aculeata L., Fabaceae, Jerusalem thornParthenium hysterophorus L., Asteraceae, partheniumParthenocissus quinquefolia (L.) Planch., Vitaceae, Virginia cree-

per #Paspalum

dilatatum Poir., Poaceae, common paspalumnotatum Flüggé, Poaceae, *2006 (PRE 1944)quadrifarium Lam., Poaceae urvillei Steud., Poaceae, tall paspalum

Passifl oracaerulea L., Passifl oraceae, blue passion fl oweredulis Sims, Passifl oraceae, purple granadillasp., Passifl oraceae suberosa L., Passifl oraceae, devil’s pumpkinsubpeltata Ortega, Passifl oraceae, granadinatripartita (Juss.) Poir. var. mollissima (Kunth) Holm-Niels. & P.Jorg.

(= P. mollissima (Kunth) L.H.Bailey), Passifl oraceae, banana pokaPennisetum

clandestinum Hochst. ex Chiov., Poaceae, Kikuyu grasspurpureum Schumach., Poaceae, Napier grasssetaceum (Forssk.) Chiov., Poaceae, fountain grasssp., Poaceaevillosum R.Br. ex Fresen., Poaceae, feathertop

Pereskia aculeata Mill., Cactaceae, pereskiaPersea americana Mill. (= P. gratissima C.F.Gaertn.), Lauraceae,

avocado pear #Persicaria lapathifolia (L.) Gray (= Polygonum lapathifolium L.),

Polygonaceae, spotted knotweedPhoenix

canariensis Hort. ex Chabaud, Arecaceae, Canary date palmdactylifera L., Arecaceae, real date palm

Phormium tenax J.R.Forst. & G.Forst., Phormiaceae, New Zealand fl ax #

Physalisperuviana L., Solanaceae, Cape gooseberryviscosa L., Solanaceae, sticky gooseberry

Phytolaccadioica L., Phytolaccaceae, belhambraicosandra L. (= P. octandra L.), Phytolaccaceae, forest inkberry

Pinuscanariensis C.Sm., Pinaceae, Canary pineelliottii Engelm., Pinaceae, slash pinehalepensis Mill., Pinaceae, Aleppo pinepatula Schiede ex Schltdl. & Cham., Pinaceae, patula pinepinaster Aiton, Pinaceae, cluster pinepinea L., Pinaceae, umbrella pineradiata D.Don, Pinaceae, radiata pineroxburghii Sarg. (= P. longifolia Roxb. ex Lamb.), Pinaceae, chir pinesp., Pinaceaetaeda L., Pinaceae, loblolly pine

Pistia stratiotes L., Araceae, water lettucePittosporum undulatum Vent., Pittosporaceae, Australian cheesewoodPityrogramma calomelanos (L.) Link, Adiantaceae, golden fernPlantago

lanceolata L., Plantaginaceae, narrow-leaved ribwortmajor L., Plantaginaceae, broad-leaved ribwortvirginica L., Plantaginaceae, dwarf plantain

Platanussp., Platanaceae #×acerifolia (Aiton) Willd. (= P. hispanica auct.), Platanaceae, London

planetree, *2004 #Plectranthus comosus Sims (= Coleus grandis Cramer)(P. barbatus

Andrews misapplied in South Africa), Lamiaceae, Abyssinian coleusPolygonum aviculare L., Polygonaceae, prostrate knotweedPolypogon monspeliensis (L.) Desf., Poaceae, beardgrassPomaderris kumeraho A.Cunn., Rhamnaceae, kumarahou #Pontederia cordata L., Pontederiaceae, pickerel weedPopulus

alba L., Salicaceae, white poplardeltoides W.Bartram ex Marshall, Salicaceae, match poplarnigra L. var. italica Münchh., Salicaceae, Lombardy poplar×canescens (Aiton) Sm., Salicaceae, grey poplar

Portulaca oleracea L., Portulacaceae, purslaneProsopis

glandulosa Torr. var. torreyana (Benson) Johnst., Fabaceae, honey mesquite

velutina Wooton, Fabaceae, velvet mesquitePrunus

armeniaca L., Rosaceae, apricotcerasoides D.Don, Rosaceae, Himalayan fl owering cherry, †Zimbabwe

# abundant locallypersica (L.) Batsch, Rosaceae, peachserotina Ehrh., Rosaceae, black cherry

Psidiumcattleianum Sabine (= P. littorale Raddi var. longipes (O.Berg)

Fosberg), Myrtaceae, strawberry guavaguajava L., Myrtaceae, guavaguineense Sw., Myrtaceae, Brazilian guavasp., Myrtaceae×durbanensis Baijnath ined., Myrtaceae, Durban guava

Pterocarya stenoptera C.DC., Juglandaceae, Chinese wing-nut #Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen &

S.M.Almeida (= P. lobata (Willd.) Ohwi), Fabaceae, kudzu vinePunica granatum L., Punicaceae, pomegranatePyracantha

angustifolia (Franch.) C.K.Schneid., Rosaceae, yellow fi rethorncoccinea M.Roem., Rosaceae, red fi rethorncrenulata (D.Don) M.Roem., Rosaceae, Himalayan fi rethornsp., Rosaceae

Pyrus sp., Rosaceae, pear tree ?#Quercus

canariensis Willd. (= Q. mirbeckii Durieu), Fagaceae, Algerian oak ?#cerris L., Fagaceae, Turkey oak ?#palustris Münchh., Fagaceae, pin oakrobur L., Fagaceae, English oaksp., Fagaceaesuber L., Fagaceae, cork oak ?#

Quisqualis indica L., Combretaceae, Rangoon creeper *2006 #Richardia

brasiliensis Gomes, Rubiaceae, tropical richardiahumistrata Steud., Rubiaceae, Peelton richardia


244 Bothalia 37,2 (2007)

Ricinus communis L., Euphorbiaceae, castor-oil plantRivina humilis L., Phytolaccaceae, bloodberryRobinia pseudoacacia L., Fabaceae, black locustRosa

multifl ora Thunb., Rosaceae, multifl ora roserubiginosa L. (= R. eglanteria L.), Rosaceae, eglantinesp., Rosaceae #?×odorata (Andrews) Sweet, Rosaceae, tea rose #

Rubuscuneifolius Pursh, Rosaceae, American bramblefl agellaris Willd., Rosaceaefruticosus L. agg., Rosaceae, European blackberry?pascuus L.H.Bailey, Rosaceae phoenicolasius Maxim., Rosaceae rosifolius Sm., Rosaceae sp., Rosaceae×proteus C.H.Stirt., Rosaceae, Bramble, Graskop/Sabie hybrid

Rumexacetosella L. subsp. pyrenaicus (Pourr. ex Lapeyr.) Akeroyd (= R.

angiocarpus auct.), Polygonaceae, sheep sorrelcrispus L., Polygonaceae, curly dockusambarensis (Dammer) Dammer (= R. nervosus Vahl var. usambarensis

Dammer), Polygonaceae, rumexSaccharum offi cinarum L., Poaceae, sugar caneSalix

babylonica L., Salicaceae, weeping willowcaprea L., Salicaceae, pussy willowfragilis L., Salicaceae, crack willow

Salsolakali L., Chenopodiaceae, common saltworttragus L. (= S. australis R.Br.), Chenopodiaceae, Russian tumble weed

Salvia tiliifolia Vahl, Lamiaceae, Lindenleaf sage, *2005 (PRE 1943)Salvinia molesta D.S.Mitch. (S. auriculata Aubl. misapplied in South

Africa), Salviniaceae, Kariba weedSambucus

canadensis L. (= S. nigra L. subsp. canadensis (L.) Bolli), Capri-foliaceae, Canadian elder

nigra L., Caprifoliaceae, European elder, *2004sp., Caprifoliaceae

Scheffl eraactinophylla (Endl.) Harms (= Brassaia actinophylla Endl.),

Araliaceae, Australian cabbage tree #arboricola (Hayata) Merr., Araliaceae, dwarf umbrella tree, *2005 #elegantissima (hort. Veitch. ex Mast.) Lowry & Frodin (= Dizygotheca

elegantissima (hort. Veitch. ex Mast.) R.Vig. Guillaumin), Araliaceae, *2005 #

Schinusmolle L., Anacardiaceae, pepper treeterebinthifolius Raddi, Anacardiaceae, Brazilian pepper tree

Schizolobium parahyba (Vell.) S. F.Blake var. parahyba (= S. excelsum Vogel), Fabaceae, parasol tree #

Schkuhria pinnata (Lam.) Kuntze ex Thell., Asteraceae, dwarf marigoldSenna

bicapsularis (L.) Roxb. (= Cassia bicapsularis L.), Fabaceae, rambling cassia

corymbosa (Lam.) H.S.Irwin & Barneby (= Cassia corymbosa Lam.), Fabaceae, autumn cassia

didymobotrya (Fresen.) H.S.Irwin & Barneby (= Cassia didymobotrya Fresen.), Fabaceae, peanut butter cassia

hirsuta (L.) H.S.Irwin & Barneby (= Cassia hirsuta L.), Fabaceae multiglandulosa (Jacq.) H.S.Irwin & Barneby (= Cassia multiglan-

dulosa Jacq., C. tomentosa L.f.), Fabaceae obtusifolia (L.) H.S.Irwin & Barneby (= Cassia obtusifolia L.), Fabaceae occidentalis (L.) Link (= Cassia occidentalis L.), Fabaceae, wild coffeependula (Willd.) H.S.Irwin & Barneby var. glabrata (Vogel) H.S.Irwin

& Barneby (= Cassia coluteoides Collad.), Fabaceaeseptemtrionalis (Viv.) H.S.Irwin & Barneby (= Cassia fl oribunda sensu

Brenan non Cav.), Fabaceae, arsenic bushsp., Fabaceae

Sesbaniabispinosa (Jacq.) W.Wight var. bispinosa (= S. aculeata Pers.), Fabaceae,

spiny sesbaniapunicea (Cav.) Benth., Fabaceae, red sesbania

Sigesbeckia orientalis L., Asteraceae, St. Paul’s wortSilybum marianum (L.) Gaertn., Asteraceae, milk thistleSisymbrium orientale L., Brassicaceae, Indian hedge mustardSisyrinchium sp., Iridaceae Solanum

betaceum Cav. (= Cyphomandra betacea (Cav.) Sendtn.), Solanaceae,

tree tomatocapsicoides All., Solanaceae, devil’s applechrysotrichum Schltdl. (= S. hispidum auctt. non Pers.), Solanaceae,

giant devil’s fi gelaeagnifolium Cav., Solanaceae, silver-leaf bitter applemauritianum Scop. (= S. auriculatum Aiton), Solanaceae, bugweedpseudocapsicum L., Solanaceae, Jerusalem cherryseaforthianum Andrews, Solanaceae, potato creepersisymbriifolium Lam., Solanaceae, dense-thorned bitter applesp., Solanaceae torvum Sw. (= S. mannii C.H.Wright), Solanaceae viarum Dunal, Solanaceae, tropical soda apple, *2006 (PRE 1962)

Sonchus oleraceus L., Asteraceae, sowthistleSophora cf. davidii (Franch.) Skeels, Fabaceae #Sorghum halepense (L.) Pers. (= S. almum Parodi), Poaceae, Johnson

grassSpartium junceum L., Fabaceae, Spanish broomSpathodea campanulata P.Beauv., Bignoniaceae, African fl ame treeSphagneticola trilobata (L.) Pruski (= Thelechitonia trilobata (L.)

H.Rob. & Cuatrec, Wedelia trilobata (L.) Hitchc.), Asteraceae, Singapore daisy

Spiraea cantoniensis Lour., Rosaceae, Cape may #Stellaria media (L.) Vill., Caryophyllaceae, chickweedStenocarpus sinuatus Endl., Proteaceae, fi rewheel tree ?#Styphnolobium japonicum (L.) Schott (= Sophora japonica L.),

Fabaceae, Japanese pagoda tree ?#Symphyotrichum subulatum (Michx.) G.L.Nesom var. squamatum

(Spreng.) S.D.Sundb. (= Aster squamatus (Spreng.) Hieron.), Asteraceae, swamp aster

Syncarpia glomulifera (Sm.) Nied. (= S. laurifolia Ten.), Myrtaceae, turpentine tree

Syzygiumcumini (L.) Skeels, Myrtaceae, jambolanjambos (L.) Alston, Myrtaceae, rose applepaniculatum Gaertn. (= Eugenia myrtifolia Sims), Myrtaceae,

Australian water pearTabebuia chrysotricha (Mart. ex DC.) Standl. (= Tecoma chrysotricha

Mart. ex DC.), Bignoniaceae, yellow trumpet tree ?#Tagetes minuta L., Asteraceae, khaki weedTamarix

chinensis Lour., Tamaricaceae, Chinese tamariskramosissima Ledeb., Tamaricaceae, pink tamarisksp., Tamaricaceae

Taraxacum offi cinale F.H.Wigg. agg., Asteraceae, common dandelionTecoma

stans (L.) Juss. ex Kunth, Bignoniaceae, yellow bellstenuifl ora (A.DC.) Fabris, Bignoniaceae, *2004

Tephrocactusarticulatus (Pfeiff.) Backeb., Cactaceae, paper-spine cholla ?aoracanthus (Lem.) Lem. (= ?Opuntia aoracantha Lemaire), Cactaceae sp., Cactaceae

Thevetia peruviana (Pers.) K.Schum. (= T. neriifolia Juss. ex Steud.), Apocynaceae, yellow oleander

Tipuana tipu (Benth.) Kuntze (= T. speciosa Benth.), Fabaceae, tipu treeTithonia

diversifolia (Hemsl.) A.Gray, Asteraceae, Mexican sunfl owerrotundifolia (Mill.) S.F.Blake, Asteraceae, red sunfl ower

Toona ciliata M.Roem. (= Cedrela toona Roxb. ex Willd.), Meliaceae, toon tree

Torilis arvensis (Huds.) Link., Apiaceae, spreading hedge-parsley Toxicodendron succedaneum (L.) Kuntze (= Rhus succedanea L.), Ana-

cardiaceae, wax treeTradescantia

zebrina hort. ex Bosse (= Zebrina pendula Schnizl.), Commelinaceae, wandering jew, *2005 #

fl uminensis Vell., Commelinaceae, wandering Jew, *2001 (Wells et al. 1986)

Tragopogon dubius Scop. (= T. major Jacq.), Asteraceae, yellow salsify Tridax procumbens L., Asteraceae, tridax daisyTriplaris americana L., Polygonaceae, triplaris ?#Triticum aestivum L., Poaceae, volunteer wheatTropaeolum majus L., Tropaeolaceae, nasturtiumUlex europaeus L., Fabaceae, European gorseUlmus

parvifolia Jacq. (= U. chinensis Pers.), Ulmaceae, Chinese elmprocera Salisb., Ulmaceae, English elm #sp., Ulmaceae #

Verbenabonariensis L., Verbenaceae, wild verbena


Bothalia 37,2 (2007) 245

Verbena (cont.)brasiliensis Vell., Verbenaceae, slender wild verbenaoffi cinalis L., Verbenaceae, European verbenarigida Spreng. var. rigida (= V. venosa Gillies & Hook.), Verbenaceae,

veined vervainVerbesina encelioides (Cav.) Benth. & Hook.f. ex A.Gray, Asteraceae,

golden crownbeardVinca major L., Apocynaceae, greater periwinkleViola

hederacea Labill. (= Erpetion reniforme Sweet), Violaceae, Australian violet, *2001 #

priceana Pollard (= V. sororia Willd.), Violaceae, confederate violet, *2005 #

Vitexagnus-castus L., Verbenaceae, lilac chastetree, *2004 (PRE 1975)

trifolia L., Verbenaceae, Indian three-leaf vitex, *2004 #sp., Vitaceae, grape #

Washingtonia sp., Arecaceae, petticoat palm ?#Wigandia urens (Ruiz & Pav.) Kunth var. caracasana (Kunth) D.N.Gibson

(= W. caracasana Kunth), Hydrophyllaceae, wigandia #Wisteria fl oribunda (Willd.) DC., Fabaceae, Japanese wisteria #

Xanthiumsp., Asteraceaespinosum L., Asteraceae, spiny cockleburstrumarium L., Asteraceae, large cocklebur

Yucca aloifolia L., Agavaceae, Spanish bayonetZinnia peruviana (L.) L. (= Z. multifl ora L.), Asteraceae, redstar zinnia

APPENDIX 6.—Characteristics of prominent invaders in study area

Plant name Family Origin GF W LC P SR VR Disp. agent Cult. use VCAcacia

cyclops Fabaceae ST (Aus.) t/s woo per ev seed bir, mam #c/b, bar sa, fy, skdealbata Fabaceae ST (Aus.) t woo per ev seed cop wat, ?ant,

?birsilc, #bar, orn

sa, fo, gr

decurrens Fabaceae ST (Aus.) t woo per ev seed cop wat, ?ant, ?bir

silc, #bar, orn

gr

longifolia Fabaceae ST (Aus.) t/s woo per ev seed wat, bir, ant #c/b, bar, orn fy, fomearnsii Fabaceae ST (Aus.) t woo per ev seed cop wat, ?ant,

?bir#silc, bar, orn

sa, fy, fo, gr, sk

melanoxylon Fabaceae ST (Aus.) t woo per ev seed suc bir, wat #silc, bar, orn

fy, fo

pycnantha Fabaceae ST (Aus.) t woo per ev seed wat, mam, ?bir

#silc, c/b, bar, orn

fy

saligna Fabaceae ST (Aus.) t/s woo per ev seed cop wat, mam, ant

silc, agrc, #c/b, bar, orn

sa, fy, fo, sk

Achyranthes aspera Amaranthaceae T (?Afr.) h her per ev/d seed mam none foAgave

americana var. america-na

Agavaceae T (Am.) s suc per ev seed suc win, hum orn, #bar, agrc

sa, nk

sisalana Agavaceae T (Am.) s suc per ev suc, bul wat, hum bar, #agrc, orn

sa

Ageratumconyzoides Asteraceae T (Am.) h her ann germ seed win #orn sa, fohoustonianum Asteraceae T (Am.) h her ann germ seed win #orn sa, fo

Argemonemexicana Papaveraceae T (Am.) h her ann germ seed wat, ?hum

(soil),?antnone sa

ochroleuca subsp. ochro-leuca

Papaveraceae T (Am.) h her ann germ seed wat, ?hum (soil),?ant

none sa

Arundo donax Poaceae NT (Med. & Asia)

g/r sem per ev rhz, div wat, hum #agrc, orn, bar

sa, fy, gr, nk, sk

Atriplexinfl ata Chenopodiaceae ST (Aus.) h sem per ev/d seed win #?agrc nk, sknummularia subsp. nummularia

Chenopodiaceae ST (Aus.) s woo/sem

per ev/d seed win #agrc, bar nk, sk

Azolla fi liculoides Azollaceae T (Am.) h her var var spore div wat, bir #orn gr, nkCaesalpinia decapetala Fabaceae Tl (Asia) s/c woo per ev seed wat, ?hum,

mam (cattle)

#bar, orn sa, fo, gr

Cardiospermumgrandifl orum Sapindaceae T (Am.) c sem per ev/d seed wat, win #orn sa, fohalicacabum Sapindaceae T (?Am.) c sem per ev/d seed wat, win #orn sa

Origin: ST, southern temperate, south of or straddling Tropic of Capricorn; NT, northern temperate, north of or straddling Tropic of Cancer; T, tropical, between or straddling Tropics of Capricorn and Cancer. GF, growth form: tree; tree/shrub; shrub; herb; herb/shrub; grass; grass/reed; climber; shrub/climber.

W, woodiness: woody; semi-woody; herbaceous; succulent.LC, life cycle: perennial, annual, variable, biennial.P, perennation: evergreen; deciduous; evergreen/deciduous; variable; germinative.SR, sexual reproduction: seed, spore.VR, vegetative reproduction: coppice; sucker; division; rhz, rhizome; bulbil; stolon; tuber; runner.Dispersal agent: wind; water; bird; mammal; human; ant.Cultivated use: # primary (= major) use; ornamental; barrier; cover/binder; agricultural crop; silvicultural crop; none.VC, vegetation category: savanna biome; fynbos biome; forest habitats; grassland biome; nama-karoo biome; succulent karoo biome.Afr., Africa; Am., America; Aus., Australia; Eur., Europe; Med., Mediterranean.


246 Bothalia 37,2 (2007)

Casuarinacunningham iana Casuarinaceae T (Aus.) t woo per ev seed wat, win orn, c/b, #bar foequisetifolia Casuarinaceae T (Pantrop.) t woo per ev seed wat, win orn, #c/b, bar fo

Cereus jamacaru Cactaceae T (Am.) t/s suc per ev seed div bir #orn, bar saCestrum laevigatum Solanaceae T (Am.) t/s woo per ev seed bir #orn, bar sa, foChromolaena odorata Asteraceae T (Am.) s woo per ev seed win #orn sa, foCinnamomum camphora Lauraceae NT (Asia) t woo per ev seed bir #orn, silc,

agrc fo

Cirsium vulgare Asteraceae NT (Eur., N Afr. & Asia)

h her bie germ seed win none gr

Daturaferox Solanaceae T (Am.) h her ann germ seed wat, ?hum

(soil), ?antnone sa, gr

innoxia Solanaceae T(Am.) h her var var seed wat, ?hum (soil), ?ant

none sa, gr

stramonium Solanaceae T (Am.) h her ann germ seed wat, ?hum (soil), ?ant

#agrc sa, gr

Eichhornia crassipes Pontederiaceae T (Am.) h her per ev seed div wat, hum #orn sa, fy, grEucalyptus

camaldulensis Myrtaceae T (Aus.) t woo per ev seed cop wat, win #silc, bar, orn, agrc

fy, fo, sk

diversicolor Myrtaceae ST (Aus.) t woo per ev seed cop win #silc, bar, agrc, orn

fy, fo

grandis Myrtaceae T (Aus.) t woo per ev seed cop win #silc, bar, orn, agrc

sa, fo, gr

Hakea sericea Proteaceae ST (Aus.) t/s woo per ev seed win orn, c/b, #bar fyHedychium

coccineum Zingiberaceae NT (Asia) h her per ev seed rhz bir, wat #orn focoronarium Zingiberaceae NT (Asia) h her per ev seed rhz bir, wat #orn fogardnerianum Zingiberaceae NT (Asia) h her per ev seed rhz bir, wat #orn fo

Ipomoeaindica Convolvulaceae T (Am.) c her per ev seed ?win, wat #orn sa, fopurpurea Convolvulaceae T (Am.) c her ann germ seed ?win, wat #orn sa, fo

Jacaranda mimosifolia Bignoniaceae T (Am.) t woo per d seed cop win #orn sa, foLantana camara Verbenaceae T (Am.) s woo per ev/d seed cop,

?runbir #orn, bar sa, fo, gr

Leptospermum laevigatum Myrtaceae ST (Aus.) t/s woo per ev seed win,wat orn, #bar, c/b fyLigustrum

japonicum Oleaceae NT (Asia) t/s woo per ev seed cop bir orn, #bar folucidum Oleaceae NT (Asia) t/s woo per ev seed cop bir orn, #bar fo

Litsea glutinosa Lauraceae T (Asia) t/s woo per ev seed ?cop bir #orn foMacfadyena unguis-cati Bignoniaceae T (Am.) c sem per ev/d seed cop, tub win #orn sa, foMelia azedarach Meliaceae T(Aus.) t woo per d seed cop bir, wat #orn sa, fo, grMorus alba Moraceae NT (Asia) t woo per d seed cop bir orn, #agrc sa, fo, grNephrolepis exaltata Davalliaceae T (Am.) h her per ev sp sto, tub win, hum #orn foNicotiana glauca Solanaceae T (Am.) t/s woo per ev seed cop win, wat,

?hum (soil)#orn sa, fy, nk, sk

Opuntiafi cus-indica Cactaceae T (Am.) t/s suc per ev seed div mam, bir #agrc, bar sa, fy, fo, gr,

nk, skrobusta Cactaceae T (Am.) ?t/s suc per ev seed div mam, bir #agrc, bar nkstricta Cactaceae T (Am.) s suc per ev seed div mam, bir #orn sa

Paraserianthes lophantha Fabaceae ST (Aus.) t/s woo per ev seed ?cop wat #orn, agrc fyPassifl ora edulis Passifl oraceae T (Am.) c her per ev seed mam, bir orn, #agrc foPennisetum clandestinum Poaceae T (Afr.) g her per ev/d seed rhz, sto ?win, hum #c/b, agrc foPereskia aculeata Cactaceae T (Am.) s/c suc per ev seed div bir, ?mam,

hum#bar, orn fo

APPENDIX 6.—Characteristics of prominent invaders in study area (cont.)

Plant name Family Origin GF W LC P SR VR Disp. agent Cult. use VC



Bothalia 37,2 (2007) 247

Pinuspatula Pinaceae T (Am.) t woo per ev seed win #silc, bar,

ornfo, gr

pinaster Pinaceae NT (Med.) t woo per ev seed win #silc, bar fy, foradiata Pinaceae NT (N Am.) t woo per ev seed win #silc, bar fy, fo

Populusalba Salicaceae NT (Eur., N

Afr. & Asia)t woo per d suc,cop wat silc, #bar,

ornsa, fo, gr, nk

deltoides Salicaceae NT (N Am.) t woo per d seed suc, cop wat, win #silc, agrc, orn

gr

nigra var. italica Salicaceae NT (Eur. & Asia)

t woo per ev/d suc, cop wat orn, #bar, c/b, agrc

gr

×canescens Salicaceae NT (Eur. & Asia)

t woo per ev/d suc, cop wat silc, #c/b, bar, orn

sa, fy, fo, gr, nk, sk

Prosopisglandulosa var. torreyana Fabaceae NT (N Am.) t/s woo per d seed cop mam, wat #agrc, orn sa, nk, skvelutina Fabaceae NT (N Am.) t/s woo per d seed cop mam, wat #agrc, orn sa, nk, sk

Prunus persica Rosaceae NT. (Asia) t woo per d seed ?cop hum #agrc, orn grPsidium guajava Myrtaceae T (Am.) t/s woo per ev seed suc, cop mam, bir,

hum#agrc, orn sa, fo

Pyracanthaangustifolia Rosaceae N Temp.

(Asia)s woo per ev seed ?cop bir orn, #bar gr

coccinea Rosaceae N Temp. (Eur. & Asia)

s woo per ev seed ?cop bir orn, #bar gr

crenulata Rosaceae NT (Asia) s woo per ev seed ?cop bir orn, #bar gr

Quercus robur Fagaceae NT (Eur. & Asia)

t woo per d seed ?cop wat, ?mam (squirrels)

#orn, agrc fy

Ricinus communis Euphorbiaceae T (Afr.) t/s woo var ev/d seed cop wat, hum #agrc, orn sa, fy, fo, skRobinia pseudoacacia Fabaceae NT (N Am.) t woo per d seed suc, cop wat, hum orn, #c/b,

bar, agrc gr

Rosa rubiginosa Rosaceae NT (Asia) s woo per d seed ?cop mam, bir #orn, bar, agrc

gr

Rubuscuneifolius Rosaceae NT (N Am.) s woo per ev/d seed suc, cop bir #agrc sa, fo, gr?pascuus Rosaceae NT (N Am.) s woo per ev/d seed suc, cop bir none sa, grfruticosus Rosaceae NT (Eur.) s woo per ev/d seed suc, cop bir #agrc sa, fy, fo, gr×proteus Rosaceae hybrid

origin (N Am. × S Afr.)

s woo per ev/d seed suc, cop bir none sa, gr

Salixbabylonica Salicaceae NT (Asia) t woo per d div wat, hum orn, #c/b,

agrcsa, fy, fo, gr, nk

fragilis Salicaceae NT (Eur. & Asia)

t woo per d div wat, hum orn, #c/b, ?agrc

gr

Salsola kali/tragus Chenopodiaceae NT (Eur. & Asia)

h her ann germ seed win none nk

Schinusmolle Anacardiaceae T (Am.) t woo per ev seed ?cop bir #orn, bar nk, skterebinthifolius Anacardiaceae T (Am.) t/s woo per ev seed ?cop bir orn, #bar fo

Senna didymobotrya Fabaceae T (Am.) t/s woo per ev seed ?cop wat, hum orn, #bar sa, foSesbania punicea Fabaceae T (Am.) t/s woo per ev/d seed wat #orn sa, fy, grSolanum

elaeagnifolium Solanaceae ST (S Am.) h/s sem per stems d

seed rhz ?bir none nk

mauritianum Solanaceae T (Am.) t/s woo per ev seed cop bir #orn sa, fy, fo, grpseudocap sicum Solanaceae T (Am.) h/s sem per ev seed ?cop bir #orn foseaforthianum Solanaceae T (Am.) c her per ev/d seed cop bir #orn sa, fo





248 Bothalia 37,2 (2007)

Tamarixchinensis Tamaricaceae NT (Asia) t/s woo per ev/d seed ?cop wat, win #?orn, c/b,

agrcnk, sk

ramosissima Tamaricaceae NT (Eur. & Asia)

t/s woo per ev/d seed cop, suc wat, win #?orn, c/b, agrc

nk, sk

Tithonia diversifolia Asteraceae T (Am.) s her var var seed ?win #orn sa, foXanthium strumarium Asteraceae T (Am.) h her ann germ seed wat, mam none sa, fo





2006-12-12

NOTICE ON CHANGE OF POLICY FOR REPRINTS OF BOTHALIA

As from the 12 December 2006, the policy on reprints of Bothalia articles has been amended as follows:

The author receives 20 reprints free. If there is more than one author, this number will have to be shared between them. The author will also receive a pdf file free on request (via the email or a CD included with the reprints): if there is more than one author, the co-author(s) should contact the first author for a copy.

The pdf file may only be used by the author in a private capacity as SANBI retains the copyright of the article and it may not be reproduced in another publication.

G. Germishuizen and Mrs B.A. Momberg: Editors, Bothaliafor Director: Research

2 Cussonia Avenue, Brummeria, PretoriaPrivate Bag X101, Pretoria 0001, RSA

Institute: Tel. (012) 8435000 Direct line: (012) 8435049 Fax (012) 804-3211Email: [email protected]

Invasive, naturalized and casual alien plants in southern Africa

Documents

Transcript of Invasive, naturalized and casual alien plants in southern Africa